EXTINCT AND VANISHING
MAMMALS
of the
OLD WORLD
'4NCIS HARPER
From the collection of the
n
m
Prelinger
v Jjibrary
t P
San Francisco, California
2006
6205 FRANKTOWN
CARSON CITY, NEVADA WTOl
QE881 Harper
H33 Extinct and vanishing
mammals of the old world
DATE DUE
FORESTA INSTITUTE
FOR
OCEAN
MOUNTAIN
STUDIES
6205 FRANKTOWN ROAD
CARSON CITY, NEVADA 89701
,."***? ;::;
KORDOFAN GIRAFFE (Giraffa camelopardalis antiquorum)
'Whenever I have watched them feeding on the tall feathery-leaved acacias, to which
they are very partial, or stalking slowly and majestically through the park-like
country they very commonly frequent, giraffes have always appeared to be amongst
the most graceful and beautiful of all wild creature*." FREDERICK C. SELOUS, 1914.
G p C f-i I- F D K G f J (E
EXTINCT AND VANISHING
MAMMALS
of the
OLD WORLD
by
FRANCIS HARPER
FORESTA INSTITUTE
FOR
OCEAN
MOUNTAIN
STUDIES
-j FKANKTCWN ROAD
CARSON CITY, NEVADA W70I
illustrations by
EARL L. POOLE
1945
SPECIAL PUBLICATION No. 12
AMERICAN COMMITTEE FOR INTERNATIONAL WILD LIFE PROTECTION
NEW YORK ZOOLOGICAL PARK, NEW YORK 60, N. Y.
35UOK -vIS
FTlhere are no words that can tell the hidden
J. spirit of the wilderness, that can reveal its
mystery, its melancholy, and its charm. There is
delight %i the hardy life of the open, in long rides
, rifle in hand, in the thrill of the fight with danger-
ous game. Apart from this, yet mingled with it,
is the strong attraction of the silent places, of the
large tropic moons, and the splendour of the new
stars; where the wanderer sees the awful glory of
sunrise and sunset in the wide waste spaces of the
earth, unworn of man, and changed only by the
slow changes of the ages from time everlasting.
THEODORE ROOSEVELT
African Game Trails
THE LORD BALTIMORE PRESS
BALTIMORE, MD., U.S.A.
FOREWORD
E1933 Dr. John C. Phillips, a founder and the first chairman
>f the American Committee for International Wild Life Protec-
tion, now in its twelfth year, was an official observer for our
Government at the meetings of the London Convention for the
Protection of the Fauna and Flora of Africa. He returned from
that conference, which concerned itself primarily with the larger
mammals of Africa, with the conviction that there was a basic need
for the compilation of our present knowledge concerning the recently
extinct and vanishing mammals, if we are to plan intelligently for
the future preservation of wild life in this fast-changing world.
This would be a pioneer job requiring the use of widely scattered
sources. Such a compilation could serve as a sound foundation for
future plans that would have to be developed to meet the ever-
increasing threats of extermination. This research could also spot-
light the species that are most threatened and reveal probable
causes of extinction that might suggest new lines of effective action
to improve their chances of survival.
For this task the American Committee engaged the services of
Dr. Francis Harper, an experienced mammalogist and a meticulous
research worker. Dr. Harper started the project in May, 1936,
and devoted more than three years to the work. The magnitude
of the undertaking proved to be much greater than originally ex-
pected, and the reasons for this are clearly set forth by the author
in the introduction to the present volume. He has spoken for the
Committee in the acknowledgments of assistance.
The American Committee takes this opportunity to repeat its
expression of gratitude to Dr. Harper for the hard work and care
that he has devoted to the preparation of this volume. We are
likewise grateful to Mr. Paul H. Oehser, editor of the United
States National Museum, for the supervision of this volume through
the press and for the preparation of the index.
This whole undertaking would not have been possible without
generous financial assistance. This has come from about 40 different
sources, including the American Philosophical Society, the Academy
of Natural Sciences of Philadelphia, the Boone and Crockett Club,
the Conservation Committee of the New York Zoological Society,
the American Wild Life Institute, and several members and mem-
VI FOREWORD
her organizations of the American Committee, as well as special
friends.
On account of its length the publication committee decided to
publish Extinct and Vanishing Mammals in two volumes. The
late Dr. Glover M. Allen, in a large measure, prepared the volume
on Extinct and Vanishing Mammals of the Western Hemisphere,
including also certain marine mammals of all the oceans. The
New World volume of more than 600 pages was published in 1942
as Special Publication No. 11 of the American Committee for
International Wild Life Protection. It was dedicated to the late
Dr. John C. Phillips.
The Committee appreciates the fact that ever-changing condi-
tions require additions and supplements to the data in these volumes
in order to bring them up to any given date. Nevertheless, keep-
ing the information current will be a small task compared with
the historical study, the verification of references, the biblio-
graphical research, and the evaluation and compilation of informa-
tion carried out by Harper and Allen in their pioneer work on the
(recently) extinct and vanishing mammals of the Old World and
the Western Hemisphere.
It is our sincere hope that these volumes may serve as a founda-
tion of information on which will be built future plans for the preser-
vation of vanishing species of mammals in their native habitats.
In many cases this may be most effectively brought about within a
framework of international cooperation such as the London Con-
vention or the Inter-American Convention. In other instances
a threatened species may be regarded as a sort of international
trust by the country under whose jurisdiction it may fall. For
example, if the Great Asiatic One-horned Rhinoceros (Rhinoceros
unicornis) should vanish from the earth (very few hundred survive
today) it would be a world calamity and not of concern merely to
the ruler of Assam who controls their last principal hide-out.
International wild-life conservation should be a concern of all
people! We must keep faith with our wild-life heritage and pre-
serve it for the wise use of generations to come!
HAROLD J. COOLIDGE, Jr.
(for the Committee)
Washington, D. C.
April 20, 1945
Publication Committee:
CHARLES M. B. CADWALADER
ALEXANDER WETMORE
HAROLD J. COOLIDGE, Jr.
CONTENTS
PAGE
FOREWORD v
INTRODUCTION
Origin, plan, methods 1
Acknowledgments 7
Factors in the progressive depletion of the Old World's mammalian
faunas 8
Australia 9
Malay Archipelago 10
Asia 11
Europe 13
Africa 15
Madagascar 16
The chronology of extinction 17
The record of extinction by families 21
Summary and conclusions 22
ACCOUNTS OF EXTINCT, VANISHING, OR THREATENED
MAMMALS
Order MARSUPIALIA : Marsupials 25
Family Dasyuridae : Dasyures, etc 25
Genus Antechinus : Broad-footed Marsupial Mice 25
Genus Phascogale: Brush-tailed Marsupial Rats 26
Genus Sminthopsis : Sminthopses 29
Genus Dasyurus : Native Cats 32
Genus Sarcophilus : Tasmanian Devil 38
Genus Thylacinus : Tasmanian Wolf 40
Family Myrmecobiidae : Marsupial Anteaters 43
Genus Myrmecobius : Numbats 43
Family Peramelidae : Bandicoots 47
Genus Perameles. Bandicoots 47
Genus Macrotis: Rabbit-bandicoots or Bilbies 51
Genus Chaeropus : Pig-footed Bandicoots 58
Family Phalangeridae : Phalangers, etc 60
Genus Tarsipes : Honey Possum 60
Genus Gymnobelideus : Leadbeater's Opossum 61
Genus Pseudocheirus : Ringtails 63
Family Phascolarctidae : Koalas 64
Genus Phascolarctos : Koalas 64
vii
Vlll EXTINCT AND VANISHING MAMMALS
Order MARSUPIALIA Continued. PAGE
Family Vombatidae : Wombats 72
Genus Vombatus : Common Wombats 72
Genus Lasiorhinus : Hairy-nosed Wombats 74
Family Macropodidae : Kangaroos, Wallabies, etc 77
Genus Bettongia : Rat-kangaroos 77
Genus Aepyprymnus : Rufous Rat-kangaroo 84
Genus Potorous : Rat-kangaroos 86
Genus Caloprymnus : Desert Rat-kangaroo 91
Genus Lagorchestes : Hare- wallabies 93
Genus Lagostrophus : Banded Wallaby 96
Genus Onychogalea : Nail-tailed Wallabies 98
Genus Petrogale : Rock-wallabies 102
Genus Thylogale: Pademelons 106
Genus Wallabia : Wallabies 114
Genus Macropus : Kangaroos 120
Order INSECTIVORA : Insectivores 122
Family Soricidae : Shrews 122
Genus Crocidura : Musk-shrews 122
Order PRIMATES : Primates 123
Family Lemuridae : Lemurs 123
Genus Microcebus : Dwarf Lemurs 123
Genus Cheirogaleus : Mouse Lemurs 126
Genus Phaner : Fork-marked Lemur 129
Genus Hapalemur: Gentle Lemurs 130
Genus Lemur: True Lemurs 132
Genus Lepilemur : Sportive Lemurs 144
Family Indriidae: Sifakas, Indri, and Avahis 146
Genus Propithecus : Sifakas 146
Genus Indri : Indri 155
Genus Avahi : Avahis 156
Family Daubentoniidae : Aye-aye 158
Genus Daubentonia : Aye-aye 158
Family Colobidae: Leaf-eating Monkeys 160
Genus Colobus : Colobus Monkeys 161
Family Pongidae : Anthropoid Apes 164
Genus Pongo : Orang-utan 164
Genus Gorilla : Gorillas 168
Genus Pan : Chimpanzees 176
Order EDENTATA : Edentates 181
Family Manidae : Pangolins 181
Genus Smutsia: Giant and South African Pangolin 181
Genus Phataginus: Three-cusped Pangolin 186
Genus Uromanis : Long-tailed Pangolin 189
CONTENTS IX
PAGE
Order RODENTIA : Rodents 190
Family Leporidae : Hares and Rabbits 190
Genus Pentalagus : Amami Hare 190
Family Castoridae: Beavers 191
Genus Castor: Beavers 191
Family Cricetidae : Hamsterlike Rodents 200
Genus Lophiomys: African Maned Rats 200
Family Muridae: Old World Rats 205
Genus Rattus: Typical Rats 205
Genus Mastacomys : Broad-toothed Rats 210
Genus Zyzomys: White-tailed Rats 211
Order CARNIVORA : Carnivores 211
Family Canidae : Wolves and Foxes 211
Genus Simenia : Simenian Foxes 212
Genus Canis: Wolves 213
Genus Nyctereutes : Raccoon-dogs 215
Family Ursidae : Bears 217
Genus Ursus : Bears 217
Family Mustelidae : Weasels, etc 232
Genus Mustek: Weasels, Minks, and Stoats 233
Genus Martes : Sables and Martens 235
Genus Gulo : Wolverines 241
Family Viverridae: Civets, Mongooses, etc 244
Genus Arctictis : Binturongs 244
Genus Fossa : Fossane 249
Family Protelidae : Aard- wolves 250
Genus Proteles : Aard- wolves 250
Family Felidae: Cats 254
Genus Cryptoprocta : Fossa 254
Genus Felis: Cats 256
Genus Lynx : Lynxes 265
Genus Caracal : Caracals 272
Genus Acinonyx : Cheetahs 274
Genus Leo : Lions 288
Genus Panthera : Leopards and Tigers 299
Order PROBOSCIDEA : Proboscideans 310
Family Elephantidae : Elephants 310
Genus Elephas: Asiatic Elephants 311
Genus Loxodonta: African Elephants 316
Order PERISSODACTYLA : Odd-toed Ungulates 322
Family Equidae: Horses, Zebras, and Asses 322
Genus Equus: Horses, Zebras, and Quagga. . . : 322
Genus Asinus: Asses . . 345
X EXTINCT AND VANISHING MAMMALS
Order PERISSODACTYLA Continued. PAGE
Family Tapiridae : Tapirs 371
Genus Acrocodia: Malay Tapir 372
Family Rhinocerotidae : Rhinoceroses 375
Genus Rhinoceros: Asiatic One-horned Rhinoceroses 375
Genus Dicerorhinus : Asiatic Two-homed Rhinoceroses 390
Genus Diceros : Black Rhinoceroses 396
Genus Ceratotherium : White Rhinoceroses 402
Order ARTIODACTYLA : Even-toed Ungulates 414
Family Hippopotamidae : Hippopotamuses 414
Genus Hippopotamus: Common Hippopotamuses 414
Genus Choeropsis: Pygmy Hippopotamus 419
Family Camelidae : Camels and Llamas 421
Genus Camelus : Camels 421
Family Tragulidae : Chevrotains 425
Genus Hyemoschus : Water Chevrotains 425
Family Moschidae : Musk Deer 427
Genus Moschus: Musk Deer 427
Family Cervidae : Deer 435
Genus Muntiacus: Muntjaks 435
Genus Rucervus : Swamp Deer 436
Genus Cervus : Red Deer and Sikas 443
Genus Elaphurus : Pere David's Deer 467
Genus Rangifer : Reindeer 469
Family Giraffidae : Giraffes and Okapi 484
Genus Giraffa : Giraffes 484
Genus Okapia : Okapi 506
Family Bovidae: Cattle, Sheep, Goats, and Antelopes 510
Genus Novibos: Cambodian Wild Ox 510
Genus Bos: Cattle 511
Genus Bibos: Gaurs and Bantengs 514
Genus Poephagus : Yaks 528
Genus Bison : Bisons 531
Genus Bubalus : Asiatic Buffaloes 538
Genus Anoa : Dwarf Buffaloes 548
Genus Syncerus : African Buffaloes 554
Genus Ovis : Sheep 557
Genus Ammotragus : Audads 600
Genus Capra : Goats and Ibexes 606
Genus Capricornis : Serows 635
Genus Alcelaphus : Hartebeests 642
Genus Damaliscus : Bontebok and allies 653
Genus Connochaetes : Gnus 659
Genus Cephalophus : Duikers 663
CONTENTS XI
Order ARTIODACTYLA Continued.
Family Bovidae Continued. PAGE
Genus Oreotragus : Klipspringers 668
Genus Nesotragus : Sunis 672
Genus Dorcatragus : Beira 674
Genus Ammodorcas : Dibatag 675
Genus Saiga : Saiga Antelope 677
Genus Gazella: Gazelles 683
Genus Aegoryx: White Oryx 690
Genus Oryx: Oryxes 693
Genus Hippotragus : Roan and Sable Antelopes and Blaauw-
bok 698
Genus Addax : Addax 711
Genus Tragelaphus : Bushbucks 716
Genus Taurotragus : Elands 722
BIBLIOGRAPHY 733
INDEX . 811
XIV EXTINCT AND VANISHING MAMMALS
FIGURE PAGE
26. Binturong: Arctictis binturong subsp. (After photograph in
Brehm and specimen in Philadelphia Zoo) 246
27. Aard-wolf: Proteles cristatus subsp 253
28. Fossa: Cryptoprocta ferox Bennett (After photograph in Brehm) .255
29. European Lynx : Lynx lynx lynx (Linnaeus) (After Standard Nat.
Hist., etc.) 266
30. Barbary Lynx: Caracal caracal algirus (Wagner) (From specimen
in Philadelphia Zoo) 273
31. Indian Cheetah: Acinonyx jubatus venaticus (Hamilton Smith). 284
32. King Cheetah: Acinonyx rex Pocock (After Pocock, 1927) 287
33. Mongolian Wild Horse: Equus przewalskii Poliakov (After photo-
graph in Brehm) 326
34. Quagga: Equus quagga Gmelin (After Standard Nat. Hist.) 335
35. Burchell's Zebra: Equus burchellii burchellii (J. E. Gray) (After
Brehm) 340
36. Mountain Zebra: Equus zebra zebra Linnaeus (After photo-
graphs in Brehm and Newnes) 343
37. Nubian Wild Ass: Asinus asinus africanus Fitzinger 346
38. Indian Wild Ass: Asinus hemionus khur (Lesson) 365
39. Malay Tapir: Acrocodia indica (Desmarest) 373
40. Great Indian Rhinoceros : Rhinoceros unicornis Linnaeus 376
41. Javan Rhinoceros: Rhinoceros sondaicus Desmarest 382
42. Northern White Rhinoceros: Ceratotherium simum cottoni
(Lydekker) (After Lang) 408
43. Pygmy Hippopotamus: Choeropsis liberiensis (Morton) 420
44. Water Chevrotain: Hyemoschus aquaticus subsp. (After Brehm) . 426
45. Schomburgk's Deer: Rucervus schomburgki Blyth 437
46. Barbary Stag : Cervus elaphus barbarus Bennett 458
47. Pere David's Deer: Elaphurus davidianus Milne Edwards 468
48. Spitsbergen Reindeer: Rangifer platyrhynchus (Vrolik) (After
Wollebaek, 1926) 480
49. Southern Giraffe: Giraffa camelopardalis capensis (Lesson) (After
Brehm, Lydekker, etc.) 504
50. Okapi: Okapia johnstoni (P. L. Sclater) 507
51. Caucasian Bison: Bison bonasus caucasicus Hilzheimer 538
52. Common Anoa: Anoa depressicornis (Hamilton Smith) 551
53. European Mouflon: Ovis musimon (Pallas) 575
54. Barbary Sheep : Ammotragus lervia subsp 600
55. Mediterranean Ibex: Capra pyrenaica hispanica Schimper (After
Lydekker and Ward) '. 613
56. Abyssinian Ibex: Capra walie Riippell (From photograph, Field
Museum) * 624
57. Markhor: Capra falconeri subsp. (After Cassell) 628
ILLUSTRATIONS XV
FIGURE PAGE
58. Sumatran Serow: Capricomis sumatraensis sumatraensis (Bech-
stein) (After Mohr, 1934) 636
59. Bubal Hartebeest: Alcelaphus buselaphus buselaphus (Pallas)
(After photograph in Brehm) 644
60. Bontebok: Damaliscus dorcas (Pallas) (After Brehm and
Lydekker) 655
61. White- tailed Gnu: Connochaetes gnou (Zimmermann) 662
62. Saiga: Saiga tatarica (Linnaeus) (After Brehm) 681
63. White Oryx: Aegoryx algazel (Oken) (From specimen in Phila-
delphia Zoo) 692
64. Blaauwbok: Hippotragus leucophaeus (Pallas) (After Daniell, in
Jardine's Naturalist's Library) 699
65. Giant Sable Antelope: Hippotragus variani Thomas 709
66. Addax: Addax nasomacidatus (Blainville) (After Brehm, etc.) .. 713
67. Nyala: Tragelaphus angasii Angas (After Sclater and Thomas,
1900) 718
THIS INVESTIGATION HAS BEEN AIDED BY A GRANT FROM THE
PENROSE FUND OF THE AMERICAN PHILOSOPHICAL SOCIETY
INTRODUCTION
ORIGIN, PLAN, METHODS
THE present work had its origin in a strongly felt need for defi-
nite information on the mammals that have become extinct
during the Christian Era, on those that are now threatened with the
same fate, on the factors contributing to the progressive depletion of
the world's mammalian faunas, and on the measures that have been
hitherto or may be hereafter undertaken for their preservation.
It consists to a large extent of an inventory of vanishing resources,
as an essential step in their conservation.
The plan and the inception of this investigation are due to the
keen interest and foresight of the late Dr. John C. Phillips, founder
and first chairman of the American Committee for International
Wild Life Protection. The work has been carried out under the
auspices of that organization and has been supported in part by
a grant from the Penrose Fund of the American Philosophical
Society. The Academy of Natural Sciences of Philadelphia was
chosen as the headquarters of the investigation, largely on account
of the very exceptional resources of its library in the literature of
natural history.
As originally projected, the investigation was to have covered the
entire world and the results were to be published in a single volume.
Owing to limitations of time, space, and available funds, as well
as the unforeseen magnitude of the task, the present volume is
restricted to the mammals of the Old World. The major part of
my work was concluded early in 1939; in only a few instances,
therefore, has it been possible to take into account the subsequently
published literature. Another volume, prepared in large part by
Dr. Glover M. Allen, late curator of mammals at the Museum of
Comparative Zoology, and published in December, 1942, deals with
the mammals of the New World and with the marine forms.
It was also hoped to include in the Introduction a general sur-
vey of conservation conditions so far as they affect mammals
in the various countries of the world. Although it has not been
possible to carry out this feature, fortunately the need for it has
1
2 EXTINCT AND VANISHING MAMMALS
been obviated in part by Brouwer's The Organisation of Nature
Protection in the Various Countries (1938) - 1
In the preparation of the accounts of the various mammals
treated, the aim has been to assemble and to present in concise form
such information as could be obtained on the following points:
Former range and numbers;
Present range and numbers (of vanishing species) ;
Date and rate of disappearance in each country (of species that have
become extinct, either locally or completely) ;
Causes of depletion or extinction, either direct or indirect;
Economic uses or importance;
Esthetic considerations;
The meiBures that have been or might be undertaken for the preservation
of each vanishing species.
The primary source of this information has been the published
literature. For this purpose, the library of the Academy of Natural
Sciences of Philadelphia has been the mainstay. In addition, I have
drawn to some extent upon the library resources of the United States
National Museum, the Museum of Comparative Zoology, the Ameri-
can Museum of Natural History, the Charles Sheldon Collection
at Yale University, and the American Philosophical Society.
An especially valuable source of information has been corres-
pondence with zoologists and conservation officials in most of the
countries of the Old World. By means of questionnaires, distributed
for the most part through the collaboration of the International
Office for the Protection of Nature in Brussels, a great mass of
fresh and largely unpublished data on the distribution, numbers,
economic status, and conservation of mammals has been assembled.
The unselfish cooperation of these contributors, on a scale per-
haps unprecedented in this field, has been an extremely helpful
and highly appreciated feature of the investigation.
Additional material and documents bearing upon the present
subject had been accumulating for some years in the office of the
American Committee for International Wild Life Protection, and
these have been utilized to considerable advantage. 2
Perhaps few zoologists have had better occasion than myself to
become impressed with the inexhaustible nature (and at the same
time the inadequacy) of the literature on systematics, distribution,
1 Special publication of the American Committee for International Wild Life
Protection, No. 9.
2 Dr. Glover M. Allen, in making use of office data of this sort in his com-
panion volume on mammals of the Western Hemisphere (1942), seems to have
been under the erroneous impression that I was responsible for gathering prac-
tically all of them, and consequently he has mentioned my name with the best
of intentions but with considerably greater frequency than the facts would
warrant. Credit for many of the data is due to sources indicated above.
INTRODUCTION O
economics, life histories, and related phases in the study of mammals.
Likewise few can become more conscious than myself of the incom-
pleteness of the present report on the points it endeavors to cover.
The chief handicap has been the sheer limitations of time, despite
unremitting labor during a period of practically three years. Sec-
ondary handicaps have been the nonavailability of certain litera-
ture, and the virtually unusable nature (to an Anglo-Saxon) of
much of the literature in the Slavic and Oriental languages.
In nearly every case a separate account has been provided for
each species or subspecies coming within the scope of the present
report. In matters of taxonomy and nomenclature I have endeav-
ored to follow the best authorities available, as exemplified in recent
monographs, catalogues, or check-lists. However, unanimity of
opinion on every detail is not to be expected of the specialists in this
field.
A really surprising amount of confusion in the nomenclature of
even some of the largest and best-known of the Old World mammals
has come to light as an incidental feature of the present investi-
gation. This seems to be due largely to lack of proper attention to
type descriptions and type localities, and to some extent to dis-
regard of the International Rules of Zoological Nomenclature. I
have attempted to straighten out some of the major nomenclatural
difficulties in two preliminary papers (Harper, 1939, 1940) , while a
few minor points, relating especially to type localities, are touched
upon in the present work.
Each account furnishes, in addition to the technical name of the
mammal under discussion, its common names in English and (if
known) in French, German, Italian, and occasionally other lan-
guages of western Europe. No attempt has been made to compile
names from unfamiliar or unwritten languages, and in only a few
exceptional cases have any been included. This statement, however,
does not apply to such native names as may have been taken over
bodily into the English or other European languages.
After the common names comes the original reference, or citation
of the type description. I have been able to verify probably 95
percent of these original references in the library of the Academy
of Natural Sciences of Philadelphia, and a few others elsewhere.
A statement of the type locality is then added in parentheses ; as far
as feasible, it is given in the form of an exact quotation from the
original description. In many cases brief supplementary or explana-
tory remarks are called for.
No attempt has been made to supply a complete list of synonyms,
and usually none whatever are cited. In certain cases, however,
where circumstances seem to render it advisable, one or more
synonyms are cited. For example, if the name considered valid and
4 EXTINCT AND VANISHING MAMMALS
adopted here happens to be less familiar than one or more that are
replaced, the latter may be cited. Or, if certain recently proposed
names are not considered valid, these may likewise be cited in
synonymy.
Persons interested in the study of mammals have frequent need
of consulting good illustrations of the various species and subspecies.
For this reason a special point has been made of supplying refer-
ences to such illustrations. While such a list of references can rarely
be exhaustive, it is believed that the hundreds of references given
here include at least a majority of the good published figures or
plates of the mammals under discussion. In the case of slightly
differentiated subspecies, an outstanding difficulty very frequently
encountered has been the determination of the particular one
figured. Even if the author or artist has provided a trinomial
designation and this is far from being a universal practice there
is always a possibility of misidentification, or of eventual refinement
of classification that will throw the identification into doubt,
unless the geographical provenance of the specimen figured is
accurately stated. The frequent disregard of this rule is the cause
of endless vexation, and it detracts seriously from the value of the
figures as zoological illustrations. Thus, in many cases the sub-
specific identity of a figure mentioned in the references is in doubt,
and some of these cases are indicated by a parenthetical query:
"subsp.?" This uncertainty of identification extends inevitably
even to some of Mr. Poole's excellent drawings that illustrate the
present text. For some are based upon "zoo" specimens, many of
which are notoriously of uncertain provenance or even represent the
hybrid offspring of different subspecies in captivity ; while others are
based upon previously published figures, themselves of somewhat
uncertain subspecific identity.
A brief description of each species or subspecies is included. In
its preparation I have aimed to utilize the type description so far
as it is at all adequate; but in many cases later and more complete
or more accurate descriptions have necessarily been drawn upon
for at least some of the characters. Constant caution is required,
however, in making use of reviews, catalogues, or monographs in
which the descriptions may be based upon specimens of unspecified
provenance. In all possible cases I have indicated the source of the
information by a direct quotation or, in the case of translation or
paraphrasing, by at least a bibliographical reference.
It may be remarked here that the entire report is documented
with such references to the fullest possible extent, not only as a
matter of simple justice to the authors of the works drawn upon,
but as an essential aid to the reader in verifying statements, and in
ascertaining what source material has been utilized on the one hand,
INTRODUCTION 5
or overlooked or disregarded on the other hand. The common
literary sins of failing to acknowledge sources of information, of
giving incomplete references, and of taking liberties with quotations,
have been scrupulously avoided as far as has lain within my power.
These matters have called for the closest possible attention in a
work that is so largely a compilation as the present one.
Perhaps no two mammalogists would agree completely on just
what species or subspecies come properly within the scope of this
report. In the first place, that scope is not completely explained in
the rather brief title chosen. With the exception of a few partially
aquatic species, such as the hippopotamuses, only land mammals are
included. The various marine and fresh-water species are dealt with
in Dr. Allen's volume (1942). A somewhat more exact but unduly
awkward title might have been Land Mammals of the Old World
that are Extinct, or Vanishing, or in Need of Special Protection.
Some of the forms included are no doubt actually increasing under
protection at the present moment but nevertheless deserve and
require the fullest possible care in order that they may continue to
survive.
It has been deemed advisable to include all African mammals
accorded protection in Schedules A and B of the London Convention
of 1933, even if subsequent investigation has shown that certain
forms are in less urgent need of close protection than was at first
supposed. On the o^her hand, the simple limitations of time and
funds have excluded a certain number of rare and more or less en-
dangered species whose status is probably more unsatisfactory than
that of a good many included species.
Finally, there are doubtless a considerable number of other mam-
mals (especially small, inconspicuous, or secretive species) that have
progressed far toward the vanishing point, or that have actually
become extinct, without their status having become known to zoolo-
gists. There is no royal road to the discovery of such a state of
affairs. Time and again extinction has taken place years in advance
of the fact coming to scientific attention. Thus, at the very best,
the present report could embody no more than a certain portion of
the current (and decidedly incomplete) knowledge on the subject.
A few words may be said here on the difficult subject of the
arrangement or sequence of the systematic groups families, genera,
species, and subspecies. The present arrangement of families is ac-
cording to Simpson (1931). Beyond this point there is apparently
no single, comprehensive, up-to-date guide to be followed. Many
recent authors of faunal lists or catalogues do not even undertake
an explanation of the sequence they adopt. For the large group of
ungulate mammals Lydekker's well-known catalogue (1913-1916)
furnishes a convenient guide in the arrangement of genera, species.
6 EXTINCT AND VANISHING MAMMALS
and subspecies. In the same way Iredale and Troughton's Australian
check-list (1934) serves for the marsupial groups. In other groups
(such as the Carnivora and the Primates) I have merely attempted
to follow general usage in so far as any such usage has been
discoverable. In the category of subspecies, the original, "nominate,"
or "typical" subspecies is introduced first, and is followed by the
others, usually in a more or less geographical sequence from north to
south or east to west.
Under each family heading a brief paragraph has been introduced,
stating the general distribution of the family, the number of genera
and species or subspecies it contains (exclusive of fossil forms) , and
the number of forms that have called for discussion in Dr. Allen's
preceding volume and in the present volume. The number so
treated varies from one in each of several families to more than a
hundred in the cattle family (Bovidae). Since no indigenous land
mammals occur in Antarctica, there is no need of further mention
of this region in the distributional statements.
The 1933 London Convention for the Protection of the Fauna and
Flora of Africa is mentioned with considerable frequency in this
volume. Since some readers may not be familiar with this Con-
vention and its far-reaching importance in the cause of international
wildlife preservation, a few words of explanation are inserted here.
The conference was called by invitation of Great Britain and was
attended by accredited representatives of the nine countries having
territories in Africa. The Convention became effective in January,
1936, when it had been ratified by five of the nine participating
governments. By January, 1940, ratification by three more countries
had taken place. Among the measures agreed upon by the Con-
vention are the establishment of national parks and nature reserves,
the regulation of traffic in animals, and the prohibition of encircling
fires and (wherever possible) of the use of poison, dazzling lights,
nets, and traps for hunting animals. The Annex to this Convention
lists, as Class A species, 17 mammals, 3 birds, and 1 plant, for
which rigid protection is agreed upon. It also lists, as Class B
species, 13 mammals and 9 birds which, although not requiring
such rigid protection, may be taken only under a special form of
license. The number of mammals so protected is actually much
larger than indicated in the above statements; for example, the
40 forms of Madagascar lemurs count as only a single item in the
list, and the same is true of the dozen subspecies of Giraffes. Further
details are set forth in Special Publications 6 and 10 of the American
Committee for International Wild Life Protection (1935 and 1940).
INTRODUCTION 7
ACKNOWLEDGMENTS
The chief burden of promoting the present investigation, in both
a spiritual and a financial sense, was magnanimously assumed in
the fir.st place by the late Dr. John C. Phillips, not merely in his
capacity as chairman of the American Committee for International
Wild Life Protection, but as a more or less personal responsibility.
The main lines of the investigation have been carried out as origi-
nally planned by him. Other members of the Committee have also
made generous contributions of funds, information, and advice.
When the magnitude of the task began to exceed all original esti-
mates, a grant from the Penrose Fund of the American Philosophical
Society provided timely aid. After Dr. Phillips's death in November,
1938, a subcommittee, consisting of Dr. Alexander Wetmore, Mr.
Charles M. B. Cadwalader, and Mr. Harold J. Coolidge, Jr., by
vigorous action found the means for completing the investigation.
I am further and particularly indebted to Mr. Cadwalader, as
director and president of the Academy of Natural Sciences of Phila-
delphia, for the provision of desk space and library facilities in
this institution. The already great resources of the Academy's
library have been considerably augmented, during and in behalf of
this investigation, by the acquisition of numerous important works
on mammals, through the efforts of Mr. Cadwalader, Mr. Brooke
Dolan, II, and Mr. George L. Harrison.
The whole-hearted cooperation of the International Office for the
Protection of Nature in Brussels, and particularly of its Secretary,
Mrs. Tordis Graim, is most gladly and gratefully acknowledged.
Mrs. Graim has generously undertaken and admirably fulfilled the
task not only of distributing questionnaires to numerous zoologists
and conservation officials in the Old World, but also of translating
and compiling the very valuable data thus obtained.
Through the courtesy of Dr. H. E. Anthony and Mr. George G.
Goodwin, of the American Museum of Natural History, lengthy
portions of indispensable works in Russian by Ognev and Nasonov
have been translated at that institution and placed in my hands.
Thereby a great deal of important information, not generally
available to non-Russian zoologists, has been incorporated in the
pages of the present work.
I must not omit to mention the patience and accommodation of
Dr. Remington Kellogg, of the United States National Museum,
during the hours I have spent in his office, consulting various works
not available in Philadelphia.
No words of mine can add to the value of the drawings produced
by the masterful strokes of Earl Poole's pen. They will be appre-
ciated by the reader not only as unusually faithful delineations of
8 EXTINCT AND VANISHING MAMMALS
the mammals they represent, but as a welcome embellishment of
the long pages of text.
The host of correspondents and collaborators, who have contrib-
uted first-hand and hitherto unpublished information of very ex-
ceptional value, and whose names will be found in proper place
on scores of the following pages, deserve the highest gratitude of
the sponsors and the compiler of this report. Without their contri-
butions the work would have been deprived of one of its most
essential features.
Finally, the most cordial thanks are due to the various authors
and publishers whose books and papers have been utilized in the
preparation of this work. It is hoped that they will be rewarded in
part, at least, by the complete acknowledgment of all items of
information so derived.
Of the following accounts of Old World species, 17, which
Dr. Francis Harper did not have opportunity to prepare on account
of taking up other investigations, were written by Glover Morrill
Allen and are subscribed with his initials. These accounts are in
large part based on the data already brought together by Dr. Harper,
to whom every credit is due for the extensive research and corres-
pondence which he undertook in order to assemble the essential
facts. The 17 accounts are: Crocidura juliginosa trichura, Christ-
mas Island Shrew; Rattus macleari, Captain Maclear's Rat; Rattus
nativitatis, Christmas Island Burrowing Rat; Colobus polykomos
and Colobus badius races, Colobus Monkeys; Pan troglodytes and
races, the Chimpanzee; Pongo pygmaeus, the Orang-utan; Hippo-
potamus amphibius and races, the Hippopotamuses; Choeropsis
liberiensis, Pygmy Hippopotamus ; Hyemoschus aquations and races,
Water Chevrotains; Cervus elaphus barbarus, North African Red
Deer; Loxodonta africana africana, South African Bush Elephant;
Diceros bicornis and races, Black Rhino ; Equus burchellii burchellii,
Burchell's Zebra; Equus zebra and race, Mountain Zebra; Equus
quagga, the Quagga; Oryx gazella and race, Gemsbok; Aegoryx
algazel, Scimitar Oryx; and Syncerus caffer caffer, Cape Buffalo.
G. M. A.
FACTORS IN THE PROGRESSIVE DEPLETION OF THE
OLD WORLD'S MAMMALIAN FAUNAS
IN the course of the present studies on the mammals that have
become extinct during the Christian Era, and on others that are
now threatened with the same fate, it has become convincingly
evident that the process of extinction is taking place at a steadily
accelerated rate. During this period of approximately 2,000 years,
INTRODUCTION 9
the world has lost, through extinction, about 106 known forms of
mammals. About 28 percent of these are subspecies of still existing
species, but the full species completely and irretrievably lost number
approximately 77.
Between A. D. 1 and 1800, about 33 mammals are more or less
definitely known to have become extinct (see list, pp. 17-18) . Each
half -century period since 1800 shows a steadily increasing rate of
extinction. The last 100 years have witnessed the passing of about
67 percent of the 106 extinct forms. In the past 50 years approxi-
mately 38 percent as many forms have been exterminated as in all
previous recorded history. At the present time more than 600 others
require consideration as vanishing or threatened forms. It is well
within the bounds of possibility that during the next hundred years
we may be extinguishing this group at the approximate rate of one
form per year.
In seeking the causes of this world-wide tragedy, it becomes ap-
parent that conditions vary widely over the different regions of
the globe, although there is a single major underlying factor nearly
throughout.
For the purposes of the present inquiry, we may here pass briefly
in review the major regions that are covered in this volume:
Australia, the Malay Archipelago, Asia, Europe, Africa, and Mada-
gascar.
AUSTRALIA
Conditions in Australia are peculiar and exceptional, owing to
the fact that its unique native mammalian fauna is predominantly
marsupial, and so lowly organized as to be quite unfitted for coping
with certain exotic and aggressive species introduced by civilized
man. The chief of these are the European Red Fox, the Domestic
Cat, the European Rabbit, the House Rats, and the House Mouse.
Further competition results from the encroachment of hosts of
sheep and cattle upon the ancestral grazing grounds of the her-
bivorous marsupials. An apparently minor predatory role is played
by the Dingo (Cams dingo), which was presumably introduced by
aboriginal man.
The Fox and the Cat (which has become feral in large numbers)
have long been active in the direct extermination of the smaller and
comparatively helpless marsupials. The Rabbit, in millions, operates
indirectly but no less effectively by overrunning the land, occu-
pying all available burrows, and depriving the herbivorous mar-
supials (even such large species as the kangaroos) of the food
necessary to their existence. The introduced rats and mice usurp
the habitats of the native species. Even sanctuaries are not proof
against such enemies as the foregoing.
10 EXTINCT AND VANISHING MAMMALS
The serious depletion of the native fauna by these agencies is
supplemented by widespread bush fires, by conversion of a vast
acreage of wild land into crop or grazing lands, by the huge fur
trade, by epizootic disease, and by the large-scale use of poisoned
bait, which takes toll of many animals besides the pests against
which it is directed.
Altogether, the situation in Australia has gotten largely beyond
human control. The rapidly growing list of extinct forms already
contains at least the following 11 :
Freckled Marsupial Mouse (Antechinus apicalis)
New South Wales Barred Bandicoot (Pemmeles jasciata)
Western Barred Bandicoot (Perameles myosura myosura)
Nalpa Bilby (Macrotis lagotis grandis)
Leadbeater's Opossum (Gymnobelideus leadbeateri)
Gaimard's Rat-kangaroo (Bettongia gaimardi)
Gilbert's Rat-kangaroo (Potorous gilbertii)
Broad-faced Rat-kangaroo (Potorous platyops)
Parma Wallaby (Thylogale parma)
Toolach Wallaby (Wallabia greyi) *
White-tailed Rat (Zyzomys argurus argurus)
Dr. W. K. Gregory, of the American Museum of Natural History,
says (1924, p. 11) : "Late in the eighteenth century, there arrived
in Australia by far the most destructive placental mammal the
world has ever seen, Homo sapiens, variety europaeus, who has
devastated the continent and is now completing the work of
destruction."
MALAY ARCHIPELAGO
Insular faunas are of extraordinary interest because of their
tendency toward endemism and because of the light they throw
upon geological history and evolutionary processes. At the same
time, by reason of the more or less strictly circumscribed nature of
their habitats, and by reason of a certain lack of adaptability or
self-defense, they are peculiarly vulnerable to attack and extermi-
nation by enemies of foreign origin. Thus the Malay Archipelago
commands the attention of the conservationist as well as of the
evolutionist. Incidentally, it was in this environment, in the fertile
mind of Alfred Russel Wallace, that one of the germs of the evolu-
tionary idea developed.
So far this region, containing the richest insular faunas of the
entire world, has fared moderately or at least comparatively well,
having lost only three mammals, all from tiny Christmas Island,
lying some 200 miles off the south coast of Java. These are a shrew
(Crocidura fuliginosa trichura) and two species of indigenous rats
(Rattus macleari and R, nativitatis) , all of which have succumbed
i A single captive remained alive in 1938 (Troughton, 1938, p. 407).
INTRODUCTION 11
to an invasion of House Rats and Domestic Cats, either through
direct attack or through some epizootic introduced by one or both
of these animals.
On the other hand, through the archipelago generally, cultivated
areas and the native population show a strong tendency to increase ;
this is especially true of the Sunda Islands and the Philippines.
Thus the native mammals are engaged in a steady retreat into the
dwindling forests.
In the Netherlands Indies many good protective measures have
been adopted. No less than 76 nature reserves have been created,
and these may be regarded as the final refuge of the native fauna.
Hunting and export of wild animals are prohibited except under
special license.
In Borneo and New Guinea the native population is less dense
than in the Sunda Islands, and there is apparently little use by the
natives of firearms that primary factor in the extermination of
wild life.
The vanishing mammals of the archipelago, for which special
concern is felt, include the following:
Orang-utan (Pongo pygmaeus)
Sumatran Elephant (Elephas maximus sumatranus)
Javan Rhinoceros (Rhinoceros sondaicus}
Sumatran Rhinoceros (Dicerorhinus sumatrensis sumatrensis)
Babirussa (Babirussa babyrussa)
Javan Banteng (Bibos sondaicus sondaicus)
Bornean Banteng (Bibos sondaicus lowi)
Tamarao or Dwarf Buffalo of Mindoro (Anoa mindorensis)
Common Anoa of Celebes (Anoa depressicornis)
Mountain Anoa of Celebes (Anoa jergusoni)
Sumatran Serow (Capricornis sumatraensis sumatraensis)
Of these, the Javan Rhin'oceros is in the most serious condition,
being reduced to perhaps two dozen individuals.
ASIA
The fauna of this greatest of the continents has been safeguarded
in part by natural conditions. Chief among these is the sparsity of
the human population over such vast areas as the taiga and the
tundra of Siberia and the deserts of Mongolia, Chinese Turkestan,
Persia, and Arabia. The great mountain masses of the Himalaya,
Tian Shan, and Altai systems, as well as numerous lesser ranges,
have also afforded a measure of protection to the mammals adapted
to these high altitudes.
A factor in the preservation of the large mammals of Afghan-
istan and Tibet has been the exclusion of all but a handful of
foreigners. India, despite its teeming population, has not exter-
12 EXTINCT AND VANISHING MAMMALS
minated a single mammal, thanks to the protective attitude toward
game assumed both by the native rulers and by the British adminis-
tration. In China, unfortunately, there seems to be little or no
thought of the conservation of wild life on the part of the great mass
of the population.
One of the most decisive factors in the accelerated depletion of
the game resources of Asia (and of other continents likewise)
during recent years has been the increasing use of modern rifles of
high power and precision. This has been especially noticeable in
Tibet, according to reports of recent explorers, and also in Arabia.
In the deserts of Iraq and Arabia pursuit of gazelles and other
animals by motor car has recently become a very serious menace to
their survival.
The Asiatic rhinoceroses, the Saiga Antelope, such large horned
ruminants as the Wapiti and other members of the deer family, and
even the lowly pangolins, have been victimized in a peculiarly
distressing way, merely because of the apparently wholly mythical
value of the horns, scales, and other parts of the body in the Chinese
pharmaceutical trade. This belief is so deeply rooted that probably
no educational campaign would be effective in staving off the
extermination of any species at the mercy of the peoples who regard
powdered rhino horn, for example, as a panacea. Even in countries
far beyond China's borders, protection of rhinoceroses and other
species in similar demand is made extraordinarily difficult by the
fabulous prices set upon them and by the incentive for poaching
under these circumstances. When the last Asiatic rhino is gone,
and the fancied benefits from its powdered horn are no longer
available, possibly then the tragic fallacy of the whole business
will dawn upon those responsible for the extermination of this
section of the world's fauna.
Of fur-bearing animals, probably the highly prized Siberian
Sables have been subjected to severest pressure, but the Soviet
Government has created several great reserves for their protection,
and has maintained a closed season on Sables over the whole terri-
tory of the USSR.
Despite the many-sided attack upon Asiatic mammals for the
sake of their meat, hides, fur, horns, scales, and even raw body
fluids that continent has exterminated to date, as far as known,
only three forms: the Japanese Wolf (Canis hodophilax), the
Syrian Wild Ass (Asinus hemionus hemippus) , and Schomburgk's
Deer (Rucervus schomburgki) .
There are a number of others, however, for which the same fate
is more or less imminent. Notable among these are the following:
Indian Cheetah (Acinonyx jubatus venaticus)
Asiatic Lion (Leo leo persicus)
INTRODUCTION 13
Przewalski's Horse (Equus przewalskii)
Transcaspian Wild Ass (Asinus hemionus finschi)
Indian Wild Ass (Asinus hemionus khur)
Javan Rhinoceros (Rhinoceros sondaicus)
Asiatic Two-horned Rhinoceroses (Dicerorhinus sumatrensis lasiotis and
D. s. niger)
Yarkand Stag (Cervus yarkandensis)
McNeill's Deer (Cervus macneilli)
White-lipped Deer (Cervus albirostris)
Malayan Gaur (Bibos gaurus hubbacki)
Gobi Argali (Ovis ammon darwini)
Semipalatinsk Argali (Ovis ammon collium)
Anadyr Bighorn (Ovis nivicola subsp.)
There are doubtless additional forms of Asiatic Wild Sheep whose
existence is seriously threatened, but information on the present
status of certain ones is scarcely sufficient to warrant a definite
statement.
EUROPE
In view of the fact that the European type of culture has
generally had such a devastating effect upon native faunas wherever
it has spread in colonies and settlements throughout the rest of the
world, it is gratifying to find that the mammalian fauna of Europe
itself has retrograded no further than it has. The chief impover-
ishment has naturally occurred in the British Isles and other densely
populated countries of Western Europe. And yet fewer Recent
mammals have been exterminated in Europe than in North America
or Australia or Africa. They seem to number only six, as follows:
European Lion (Leo leo subsp.)
European Wild Horse (Equus caballus subsp.)
Aurochs (Bos primigenius)
Caucasian Bison (Bison bonasus caucasicus)
Pyrenean Ibex (Capra pyrenaica pyrenaica)
Portuguese Ibex (Capra pyrenaica lusitanica)
The retrogression of the European fauna has no doubt been due
in the first place to the widespread clearing of forests and their
replacement by lands devoted to habitations, transportation systems,
crops, or grazing. Hunting, however, has constituted the most im-
portant part of the direct human pressure upon the wild animals.
While this sort of pressure began to be felt ages ago, it was primarily
the invention and improvement of firearms that enabled man to
proceed with ever-increasing rapidity on his course of extermination.
Species of comparatively large size, furnishing valuable meat and
hides, have been the prinicipal sufferers. Thus four of the six
extinct European mammals are members of the cattle family
(Bovidae).
14 EXTINCT AND VANISHING MAMMALS
In Europe, as contrasted with the United States, there is a far
greater proportion of closely guarded private estates, and hunting
of large game is chiefly restricted to the wealthy few. This con-
dition of affairs has*resulted in a much slower rate of extermination
than in the United States, despite the large number of national
parks and wild-life refuges in this country. Furthermore, the
European attitude appears much more tolerant toward such preda-
tory animals as Wolves and Brown Bears, which have been able
to survive so far in such countries as Spain, France, Italy, Yugo-
slavia, Greece, Bulgaria, Rumania, Czechoslovakia, Poland, the
Baltic States, Russia, and Scandinavia. Americans have been more
ruthless in exterminating, or attempting to exterminate, any preda-
tory animal conflicting, or presumed to conflict, with human interests.
Unfortunately, the American method of dealing with predators by
means of poison has attained a certain vogue in Bulgaria.
A few of the more important vanishing mammals of Europe may
be mentioned here. The Brown Bear (Ursus arctos) and the Wolf
(Cam's lupus) are probably doomed to disappear almost entirely
from Western Europe, although they will long survive in Russia
and Siberia. The European Wildcat (Felis silvestris silvestris) has
become extremely scarce in general; perhaps its greatest danger
lies in extinction by dilution through interbreeding with feral
Domestic Cats. The insular Wildcats (Cretan, Sardinian, Corsican,
and British Felis agrius, F. sarda, F. reyi, and F. silvestris
grampia) are probably endangered in like manner. The European
Beaver (Castor fiber), persecuted for its fur, remains in only a few
isolated colonies. There is some doubt as to whether any repre-
sentatives of the Finland Reindeer (Rangifer tarandus fennicus)
and the Novaya Zemlya Reindeer (R. t. pearsoni) still survive;
the animal of Novaya Zemlya has fallen victim to visiting
ships' crews and to Samoyed immigrants. While the stock of the
Lithuanian Bison (Bison bonasus bonasus) is greatly reduced, and
while there has been considerable mixture in captivity with the
Caucasian Bison (B. b. caucasicus) and with the American Bison
(Bison bison bison), energetic protection in sanctuaries assured
its survival up to 1939, at least. Two of the four races of the
Spanish Ibex (Capra pyrenaica) have been exterminated by exces-
sive hunting, and the fate of those remaining has become uncertain
during recent events in Spain. The Cyprian Mouflon (Ovis ophion
ophion) has become reduced to a precariously small stock.
The British Isles have long since lost the Brown Bear, the Wolf,
the Beaver, the Wild Boar (Sus scrofa) , and the Reindeer (Rangifer
tarandus) . No doubt insularity has here played a part in the early
disappearance of these mammals.
INTRODUCTION 15
AFRICA
As long as the African Continent was occupied by primitive
savages, without modern weapons, animal life was, in a large
sense, in a virtual state of equilibrium. When European settle-
ment began, and firearms were introduced, the death knell of a
very considerable proportion of the population of large mammals
was sounded. Thus the Atlas Bear (Ursus crowtheri) , the Barbary
and the Cape Lions (Leo leo leo and L. I. melanochaitus) , the
Quagga (Hippotigris quagga) , Burchell's Zebra (Hippotigris bur-
chellii burchellii), the Bubal Hartebeest (Alcelaphus busclaphus
buselaphus) , the Rufous Gazelle (Gazella rufina) , and the Blaauw-
bok (Hippotragus leucophaeics) have departed finally and completely
from the African scene. The typical subspecies of the Cape Harte-
beest (Alcelaphus caama) may also be extinct, but imperfect knowl-
edge of its distribution precludes a definite statement. A long
time previously the Algerian Wild Ass (Asinus atlanticus) became
extinct, from unknown causes. These losses by extinction are
divided almost equally between South Africa the region most
thoroughly settled by Europeans and the Barbary States, where
the well-armed Moors long held sway.
A century ago the Boer hide-hunters decimated the remarkable
antelope and zebra fauna of South Africa. In the last half-century,
firearms in the hands of improvident and short-sighted natives have
wrought extremely serious havoc among the dwindling herds of
African game in general. As intertribal warfare has practically
ceased, and as the benefits of modern medicine and sanitation have
penetrated far into the jungles and deserts, the native populations
have increased, and their demands for a meat diet have decimated
the game. Encircling fires, a method of hunting practiced on a
fairly large scale in the savanna regions, have been extremely
destructive, even in the absence of firearms. Professional hunters
in the employ of great industrial enterprises, as in various parts of
the Belgian Congo, have simply wiped out the antelopes over large
areas. Hasty and probably ill-considered campaigns for the control
of the tsetse fly have too often resulted in hecatombs of the large
game mammals. In recent years the animals of the desert, such as
Oryx and Gazelles, have become subject to attack from motor cars.
In South Africa the Bontebok (Damaliscus dorcas) , the Blesbok
(Damaliscus phillipsi) , and the White-tailed Gnu (Connochaetes
gnou) no longer roam the free veldt, but have become restricted to
enclosed farms and preserves. A remnant of the Cape Mountain
Zebra (Hippotigris zebra zebra) was preserved at the eleventh hour.
16 EXTINCT AND VANISHING MAMMALS
Among other vanishing or threatened African mammals, the fol-
lowing may be mentioned in particular:
Barbary Lynx (Caracal caracal algirus)
South African Bush Elephant (Loxodonta africana ajricand)
African Manatee (Trichechus senegalensis)
Nubian Wild Ass (Asinus asinus africanus)
Somali Wild Ass (Asinus asinus somaliensis)
Southern White Rhinoceros (Ceratotherium simum simum)
Northern White Rhinoceros (Ceratotherium simum cottoni)
Pygmy Hippopotamus (Choeropsis liberiensis)
Barbary Stag (Cervus elaphus barbarus)
Congo Giraffe (Giraffa camelopardalis congoensis)
Nigerian Giraffe (Giraffa camelopardalis peralta)
Angola Giraffe (Giraffa camelopardalis angolensis)
Southern Giraffe (Giraffa camelopardalis capensis)
Okapi (Okapia johnstoni)
Cape Buffalo (Syncerus caffer cafjer, here restricted to the South African
animal)
Egyptian Arui (Ammotragus lervia ornata)
Libyan Arui (Ammotragus lervia fassini)
Nubian Ibex (Capra nubiana nubiana)
Abyssinian Ibex (Capra walie)
Cuvier's Gazelle (Gazella cuvien)
Slender-horned Gazelle (Gazella leptoceros)
Mhorr Gazelle (Gazella dama mhorr)
White Oryx (Aegoryx algazel)
Giant Sable Antelope (Hippotragus variant)
Addax (Addax nasomaculatus)
Nyala (Tragelaphus angasii)
Mountain Nyala (Tragelaphus buxtoni)
Senegambian Giant Eland (Taurotragus derbianus derbianus)
Congo Giant Eland (Taurotragus derbianus congolanus)
A very considerable number of game reserves have been estab-
lished in various parts of Africa, and there should be a great many
more of them, effectively supervised. Herein lies the chief hope for
the survival of many of the larger African mammals.
MADAGASCAR
The mammalian fauna of this great island is particularly note-
worthy for its very high degree of endemism and for the prepon-
derance of lemurs. Madagascar and its outliers boast no less than
three families and forty species and subspecies of lemurs, not one
of which extends to the African mainland. Fortunately a fair pro-
portion of these remain more or less common, being protected
from persecution by native superstition. However, one species, the
Hairy-eared Mouse Lemur (Cheirogaleus trichotis) , is apparently
extinct. The following seem to exist in very small numbers, and
INTRODUCTION 17
should be safeguarded by every possible means from further de-
crease:
Coquerel's Dwarf Lemur (Microcebus coquereli)
Crossley's Mouse Lemur (C heirogaleus major crossleyi)
Gray Lemur (Hapalemur griseus griseus)
Broad-nosed Gentle Lemur (Hapalemur simus)
Diademed Sifaka (Propithecus diadema diadema)
Major's Sifaka (Propithecus verreauxi majori)
Aye-aye (Daubentonia madagascariensm)
Two peculiar carnivores, the Fossane (Fossa fossa) and the Fossa
(Cryptoprocta ferox) , are also endemic in Madagascar. The former
is accorded protection under Schedule A of the London Convention
of 1933, and probably the latter is almost equally deserving of con-
sideration.
Perhaps the greatest danger to mammalian life in Madagascar is
the steady reduction of the forest areas through burning and clear-
ing by the natives. It is highly important from the point of view
of conservation that this process should be halted.
THE CHRONOLOGY OF EXTINCTION
It may be of interest to the historian of mammalogy to list the
extinct forms here in some sort of chronological order. They will be
arranged chiefly by half-century periods and by regions within those
periods; but those forms that passed out of existence prior to 1800
will be placed in a single group. It should be borne in mind that
in most cases the date of extinction can be only roughly indicated.
For this reason the sequence within the regional half-century groups
will be systematic rather than chronological. In some cases, how-
ever, it is possible to add a more approximate date of extinction
after the name of the species or subspecies. Certain cases of probable
but unproved extinction are indicated by a question mark.
Years 1-1800 (33 forms) :
EUROPE
European Lion (Leo leo subsp.), 80-100
European Wild Horse (Equus caballvs subsp.)
Aurochs (Bos primigenius) , 1627
AFRICA
Algerian Wild Ass (Asinus atlanticus)
Blaauwbok (Hippotragus leucophaeus] , 1800
2
18 EXTINCT AND VANISHING MAMMALS
WEST INDIES 1
Four Antillean insectivores (Nesophontes edithae; N. micrus; N.
longirostris ; N. zamicrus)
Lesser Falcate-winged Bat (Phyllops veins]
Cuban Yellow Bat (Natalus primus)
Smaller Puerto Rican Ground Sloth (Acratocnus odontrigonus)
Larger Puerto Rican Ground Sloth (Acratocnus major)
Smaller Hispaniolan Ground^loth ( Acratocnus (?) comes)
Larger Hispaniolan Ground cloth (Parocnus serus)
Barbuda Musk-rat (Megalomys audreyae)
Hispaniolan Spiny Rat two species (Brotomys voratus; B. contractus)
Cuban Short-tailed Hutia (Geocapromys columbianus)
Crooked Island Hutia (Geocapromys ingrahami irrectus)
Great Abaco Hutia (Geocapromys ingrahami abaconis)
Haitian Hexolobodon (Hexolobodon phenax)
Least Hispaniolan Hutia (Plagiodontia spelaeum)
Puerto Rican Isolobodon (Isolobodon portoricensis)
Haitian Isolobodon (Isolobodon levir)
Narrow-toothed Hutia (Aphaetreus montanus)
Two agoutilike rodents (Heteropsomys insularis; Homopsomys antil-
lensis)
A Puerto Rican hystricomorph (Heptaxodon bidens)
"Quemi" of Oviedo (Quemisia gravis), about 1550?
A Puerto Rican giant rodent (Elasmodontomys obliquus)
SOUTH AMERICA
Patagonian Giant Ground Sloth (Grypotherium listai)
OCEANS
Steller's Sea-cow (Hydrodamalis gigas), 1768
Years 1801-1850 (2 forms) :
NORTH AMERICA
Eastern Bison (Bison bison pennsylvanicus) , 1825
WEST INDIES
Hispaniolan Hutia (Plagiodontia aedium)
Years 1851-1900 (31 forms) :
AUSTRALIA
Gilbert's Rat-kangaroo (Potorous gilbertii)
i Possibly the extinction of some of the forms listed under this heading, known
from bones found in cavern deposits, may have occurred more than 2,000 years
ago. They are recognized, however, as pertaining to the Recent fauna. A. W.
INTRODUCTION 19
EUROPE
Portuguese Ibex (Capra pyrenaica lusitanica) , about 1892
AFRICA
Atlas Bear ( Ursus crowtheri)
Cape Lion (Leo leo melanochaitus) , about 1865
Quagga (Hippotigris quagga), about 1878
MADAGASCAR
Hairy-eared Mouse Lemur (Cheirogaleus trichotis)
NORTH AMERICA
Gull Island Meadow Mouse (Microtus pennsylvanicus nesophilus),
1890's
Plains Grizzly (Ursus horribUis horribUis)
California Coast Grizzly (Ursus calif ornicu*) , about 1886
Sacramento Grizzly (Ursus colusus), about 1862
Navajo Grizzly ( Ursus texensis navaho)
Sonora Grizzly (Ursus kennerleyi)
Mendocino Grizzly (Ursus mendocinensis) , about 1875
New Mexico Grizzly (Ursus horriaeus)
Sea Mink (Mustela macrodon), about 1880
Eastern Wapiti (Cervus canadensis canadensis), about 1885
Oregon Bison (Bison bison oregonus), about 1850's
WEST INDIES
?Two Antillean insectivores (Nesophontes paramicrus; N. hypomicrus)
?Puerto Rican Long-nosed Bat (Monophyllus frater)
? Jamaican Long-tongued Bat (Reithronycteris aphylla)
A Puerto Rican bat (Stenoderma rufum)
?Puerto Rican Long-tongued Bat (PhyUonycteris major)
?Haitian Long-tongued Bat (PhyUonycteris obtusa)
Jamaican Rice Rat (Oryzomys antUlarum), about 1880's
St. Vincent Rice Rat (Oryzomys victus), about 1897?
Santa Lucia Musk-rat (Megalomys luciae)
Larger Cuban Spiny Rat (Boromys offella)
Lesser Cuban Spiny Rat (Boromys torrei)
FALKLAND ISLANDS
Antarctic Wolf (Dusicyon australis), 1876
GALAPAGOS ISLANDS
Chatham Island Rice Rat (Oryzomys galapagoensis)
20 EXTINCT AND VANISHING MAMMALS
Years 1901-1944 (40 forms) :
AUSTRALIA
Freckled Marsupial Mouse (Antechinus apicalis)
New South Wales Barred Bandicoot (Perameles jasciatd)
Western Barred Bandicoot (Perameles myosura myosura)
Nalpa Bilby (Macrotis lagotis grandis)
Leadbeater's Opossum (Gymnobelideus leadbeateri)
Gaimard's Rat-kangaroo (Bettongia gaimardi)
Broad-faced Rat-kangaroo (Potorous platyops)
Parma Wallaby (Thylogale parma)
Toolach Wallaby (WaUabia greyi)
White-tailed Rat (Zyzomys argurus ar gurus)
MALAY ARCHIPELAGO
Christmas Island Shrew (Crocidura juliginosa trichura), about 1904
Maclear's Rat (Rattus macleari), about 1904
Bulldog Rat (Rattus nativitatis) , about 1904
ASIA
Japanese Wolf (Canis hodophttax)
Syrian Wild Ass (Asinus hemionus hemippus), about 1927
Schomburgk's Deer (Rucervus schomburgki) , 1930's
EUROPE
Caucasian Bison (Bison bonasus caucasicus), 1930's
Pyrenean Ibex (Capra pyrenaica pyrenaica), 1910's
AFRICA
Barbary Lion (Leo leo leo), 1922
Burchell's Zebra (Hippotigris burchellii burchellii)
Bubal Hartebeest (Alcelaphus buselaphus buselaphus), 1920's?
Rufous Gazelle (Gazella rufina), 1920's?
NORTH AMERICA
Long-eared Kit Fox (Vulpes macrotis macrotis), 1900's
Newfoundland Wolf (Canis lupus beothucus), 1910's
Florida Wolf (Canis niger niger), 1920's
Tejon Grizzly (Ursus tularensis), 1916
Texas Grizzly (Ursus texensis texensis), 1910's?
?Mount Taylor Grizzly (Ursus perturbans)
Black Hills Grizzly (Ursus rogersi bisonophagus)
?Lillooet Grizzly (Ursus pervagor)
?Klamath Grizzly (Ursus klamathensis)
Southern California Grizzly (Ursus magister), 1908
INTRODUCTION 21
?Apache Grizzly (Ursus apache)
Henshaw's Grizzly (Ursus henshawi), 1920's
Eastern Cougar (Felis concolor couguar]
Arizona Wapiti (Cervus canadensis merriami), 1906
Badlands Bighorn (Ovis canadensis auduboni), 1900's?
WEST INDIES
Cuban Solenodon (Solenodon cubanus), about 1910
Martinique Musk-rat (Megalomys desmarestii), 1902
GALAPAGOS ISLANDS
James Island Rice Rat (Nesoryzomys swarthi)
This record shows a steadily accelerated rate of extinction in
each of the last three half-century periods. About 38 percent of
the losses have been sustained since 1900. This indicates how diffi-
cult is the task of preserving native faunas in the present era of
intensive modern invention and industrial expansion.
THE RECORD OF EXTINCTION BY FAMILIES
The following record indicates how these losses by extinction are
divided among the various mammalian families:
Bears (Ursidae), 17
Spiny rats and their relatives (Echimyidae), 15
Cattle, sheep, goats, and antelopes (Bovidae), 10
Hamsterlike rodents (Cricetidae), 8
Antillean insectivores (Nesophontidae), 6
Leaf -nosed bats (Phyllostomidae), 6
Kangaroos and their relatives (Macropodidae), 5
Wolves and foxes (Canidae), 5
Horses, zebras, and asses (Equidae), 5
Ground sloths (Megalonychidae), 4
Cats (Felidae), 4
Bandicoots (Peramelidae), 3
Old World rats (Muridae), 3
Deer (Cervidae), 3
Giant rats (Dinomyidae), 2
Dasyures and their relatives (Dasyuridae), 1
Phalangers and their relatives (Phalangeridae), 1
Solenodons (Solenodontidae), 1
Shrews (Soricidae), 1
Long-legged bats (Natalidae), 1
Lemurs (Lemuridae), 1
Giant ground sloths (Megatheriidae), 1
Heptaxodon (Heptaxodontidae), 1
Weasels and their relatives (Mustelidae), 1
Steller's Sea-cow (Hydrodamalidae), 1
There is the clearest sort of significance in the losses sustained
by the larger predatory mammals as a group (Ursidae, Canidae,
22 EXTINCT AND VANISHING MAMMALS
and Felidae) , because of their competition with man for food in the
shape of the ungulate mammals, both wild and domesticated, such
as cattle, sheep, goats, antelopes, horses, asses, swine, and deer.
In the case of such formidable carnivores as wolves, bears, lions,
tigers, and leopards, the matter of outright self-defense on man's
part may also be involved. Moreover, it is natural that the large
game species of the cattle and deer families, which require extensive
feeding grounds and are eagerly sought by mankind for food,
should have suffered some of the principal losses.
SUMMARY AND CONCLUSIONS
During the past 2,000 years the world has lost, through extinc-
tion, about 106 forms (species or subspecies) of mammals. They
are distributed by regions as follows: Australia, 11; Malay Archi-
pelago, 3; Asia, 3; Europe, 6; Africa, 9; Madagascar, 1; North
America, 27; West Indies, 41; South America, 1; Falkland Is-
lands, 1; Galapagos Islands, 2; oceans, 1. Approximately 67 per-
cent of these losses have occurred during the past century, and 38
percent during the past half-century. Thus the rate of extinction
is being steadily accelerated.
In addition to the mammals already extinct, more than 600 others
require consideration as vanishing or threatened forms.
Insular faunas, partly by reason of their circumscribed nature
and partly by reason of a certain lack of adaptability or self-
defense, are particularly vulnerable to attack or competition by man
and by certain mammalian pests introduced by him. There may be
a further reason for the decadence of insular faunas in some cases,
such as that of the West Indies, in the virtually total lack of native
mammalian predators; these would doubtless have played a bene-
ficial role by eliminating the less fit individuals, and thereby con-
tributing to the survival of the fittest individuals, among the species
preyed upon.
In general, it is fairly obvious that species of restricted distribu-
tion and specialized habits have less chance of survival than those
of wide distribution and generalized habits.
The primary factor in the depletion of the world's mammalian
faunas is civilized man, operating either directly through excessive
hunting and poisoning, or indirectly through invading or destroying
natural habitats, placing firearms in the hands of primitive peoples,
or subjecting the primitive faunas of Australia and of various
islands to the introduction of aggressive foreign mammals, including
fox, mongoose, cat, rat, mouse, and rabbit. Except in the West
Indies, comparatively few species seem to have died out within the
past 2,000 years from natural causes, such as evolutionary senility,
disease, or climatic change.
INTRODUCTION 23
The chief hope for the survival of the larger mammals of the
world lies in the establishment and maintenance of a sufficient num-
ber of sanctuaries. This will avail in most parts of the world, but
the matter is not so simple in Australia. Unless sanctuaries in that
country can be surrounded with fences that are proof against foxes,
rabbits, cats, and house rats, even they will not avail for many of
the smaller Australian mammals. So perhaps the darkest picture
today, as far as the future of mammals is concerned, is to be found in
Australia, where many of the primitive native species cannot stand
up against the highly organized introduced pests, and where condi-
tions have gotten largely beyond human control.
ACCOUNTS OF EXTINCT, VANISHING, OR
THREATENED MAMMALS
Order MARSUPIALIA: Marsupials
Family DASYURIDAE: Dasyures, etc.
This family, consisting of about 13 genera, is limited to Australia,
Tasmania, New Guinea, and certain smaller neighboring islands.
Of approximately 75 known forms, 14 call for discussion in the
present work.
Freckled Marsupial Mouse
ANTECHINUS APICALIS (J. E. Gray)
Phascogale apicalis J. E. Gray, Ann. Mag. Nat. Hist., ser. 1, vol. 9, p. 518, 1842.
("Doubtless from Australasia" =. South-Western Australia, fide Iredale and
Troughton, 1934, p. 6.)
FIGS.: Gould, 1863, vol. 1, pi. 39; Cabrera, 1919, pi. 5, fig. 2.
This marsupial of Western Australia, little larger than a mouse,
does not seem to have been collected for more than 30 years and is
probably extinct.
The general color is freckled reddish gray ; eye ring whitish ; under
parts dull white or yellowish; pouch hairs dark rufous; front and
outside of forearm rufous; rest of outer surface of limbs dull gray;
ears short; tail tapering, variegated like back, and tipped with black.
Head and body, 111-120 mm.; tail, 85-89 mm. (Thomas, 1888,
pp. 277-278.)
Very little information concerning this species is on record, and
some of that is conflicting. Gould evidently gave it much too wide
a range in stating (1863, p. 46) that it "is very generally distributed
over every part of the colony of Western Australia." His actual
records are from the vicinity of Moore's River, Perth, and King
George's Sound.
Thomas (1888, p. 278) records specimens from Albany and Vic-
toria Plains, Western Australia, and even from Queensland.
Shortridge states (1910, p. 840; map, p. 842) that it is "confined
to the forest districts of the South-W T est, where it is apparently a
rare species." He records three specimens from Albany in the Perth
Museum.
25
26 EXTINCT AND VANISHING MAMMALS
On the other hand, Glauert states (1933, p. 19) that the species
is not represented in the West Australian Museum at Perth and is
now probably extinct.
Furthermore, despite the various specimens recorded by Gould,
Krefft, Thomas, and Shortridge, E. Le G. Troughton writes (in litt.,
April 16, 1937) that "this species is not represented in the Perth
Museum, and is probably known only from the type in the British
Museum, and one in the Australian Museum. Apparently extinct or
represented by small colonies only." Iredale and Troughton
(1934, p. 6) limit the range to "South -Western Australia," omitting
Queensland.
The species was probably either neutral or beneficial in its habits,
for Gilbert (in Gould, 1863, p. 46) found the remains of insects in
the stomachs he examined.
No particular reason for its extinction seems to have been sug-
gested, but the generally adverse conditions now facing the smaller
marsupials of Australia are doubtless sufficient to account for it.
Large Brush-tailed Phascogale; Brush-tailed Pouched Mouse
PHASCOGALE TAPOATAFA TAPOATAFA (Meyer)
Viverra tapoataja Meyer, Zool. Entdeck., p. 28, 1793. (Based upon "The
Tapoa Tafa" of White, Jour. Voy. New South Wales, p. 281, pi. 58, 1790;
type locality, Sydney, New South Wales.)
SYNONYM: Didelphis penicillata Shaw (1800).
FIGS.: Waterhouse, 1841, pi. 8; Gould, 1845, vol. 1, pi. 31; Lydekker, 1894,
pi. 28; Jones, 1923, p. 99, fig. 60; Le Souef and Burrell, 1926, fig. 93;
Fleay, 1934, pis. 19, 20.
Though very considerably reduced in numbers, this animal still
maintains itself in various localities through its wide range over
the southern parts of Australia.
Form stout and strong; general color finely grizzled pale gray;
muzzle with indistinct darker stripe; ears very large, thin, nearly
naked; under parts white or pale gray; pouch hairs dull rufous,
tipped with white; terminal three-fifths of tail with a thick black
brush. Head and body, 240 mm.; tail, 225 mm. (Thomas, 1888,
pp. 295-296.)
The general range is "southern Australia, from south Queensland
to Western Australia" (Iredale and Troughton, 1934, p. 7) .
Though once a familiar animal to settlers whose homes were in the more
wooded districts, P. penicillata is unknown to the rising generation of country
people. ... It seems astonishing that so small an animal could ever have
been a real menace to the poultry run of the settler, and yet it is credited
with being a determined slayer of chickens, and one which killed not merely
to appease its appetite. Many of the older residents in South Australia have
caught the animal red-handed, and as with the Native Cat, it seems a re-
ORDER MARSUPIALIA: MARSUPIALS 27
markable thing that so well equipped a carnivore should have been reduced
to a condition bordering on extinction in so comparatively short a time.
What its range within the State may have been is difficult to determine.
It was not met with by the Horn Expedition, but an animal which answers
very much to its description, but of which no specimen is available, apparently
exists over a wide area in the Centre. By the South Australian Murray River
natives it was well known under the name of "Pundi" but it has not been
seen in their district for very many years. (Jones, 1923, p. 101.)
Shortridge comments (1910, p. 839; map, p. 841) on its status
in Western Australia as follows:
"Although not plentiful this species seems to have a more general
range in South-Western Australia than the smaller Phascogales.
"Doubtfully recorded from as far inland as Kalgoorlie, where it
would probably be only a straggler. . . .
"Occasionally frequenting the neighbourhood of farms, where
according to natives they come after mice."
According to Glauert (1933, p. 19), it occurs in the southwest of
Western Australia, from Fremantle to the south coast and inland
to Merredin. Twenty specimens had been received at the Perth
Museum in the preceding five years.
For many years since the ravages of disease during the years 1898-1900,
that agile and courageous little killer, "the brush-tailed rat" of the bushman,
has been very scarce in the majority of its old haunts in Victoria and New
South Wales. . . .
The black "bottle-brush" tail and coat of 'possum-grey fur, combined with
the amazingly agile movements of this lithe rat-sized marsupial, at once excite
admiration. However, few people have enjoyed the spectacle of the nocturnal
and arboreal creature making its lightning movements up and down the
Eucalypt trunks "corkscrewing" round the boles to elude observation, or
else bounding lightly, like a squirrel, from tree to tree. (Fleay, 1934, p. 89.)
In Victoria, according to C. W. Brazenor (in litt., March 3, 1937) ,
the animal is "holding its own and common in timbered country."
Le Souef and Burrell (1926, pp. 333-336) give the following
account: "Some species, notably the brush -tailed and the lesser
brush-tailed phascogales, are now rather scarce over the greater
part of their range, having been greatly reduced by disease, which
swept off large numbers of native animals in 1898-9-1900. Cats
have also been very destructive. . . .
"This species is more carnivorous than most members of the
family. Moreover, it is very useful, in that it seems especially to
catch rats and mice. There are instances of it following up plagues
of these rodents and doing a good deal toward thinning them out."
E. Le G. Trough ton (in litt., April 16, 1937) regards it as an
active and resourceful species, whose survival is apparently assured,
at least in the mountainous regions of its range.
[A northern subspecies (P. t. pirata Thomas, 1904; type locality,
"South Alligator River," Northern Australia) ranges across the
28 EXTINCT AND VANISHING MAMMALS
northern part of the continent, from the Dawson Valley, Queens-
land, to the Kimberley Division of Western Australia. It "appears
to be very rare on the Dawson but still has a good hold on the
wetter coastal country of the Fitzroy" (Finlayson, 1934, p. 226).
It is reported as numerous in Arnhem Land, Northern Australia
(Le Souef and Burrell, 1926, p. 336) . Its range lies largely outside
that of the introduced fox, and its chances of survival are probably
better than those of the southern subspecies.]
Red-tailed Phascogale; Lesser Brush-tailed Pouched Mouse
PHASCOGALE CALURA Gould
Phascogale calurus Gould, Proc. Zool. Soc. London 1844, p. 104, 1844. ("In
the interior of Western Australia" = the Military Station on Williams
River, fide Gould, 1863, vol. 1, p. 39.)
FIGS.: Gould, 1863, vol. 1, pi. 32; Waterhouse, 1846, pi. 14, fig. 2.
The Red-tailed Phascogale, of South, Central, and Western Aus-
tralia, is so rare that few more than a dozen specimens seem to have
been placed on record. It is evidently a vanishing species.
General color ashy gray; under parts creamy white; ears large,
nearly naked except at base, where there are some yellow hairs;
basal half of tail rusty red above, black below; terminal half bushy,
black (Gould, 1844, p. 105) . Head and body, 125 mm.; tail, 147 mm.
(Thomas, 1888, p. 297.)
Shortridge writes (1910, p. 839; map, p. 840) : "Very rare, seem-
ing hitherto to have been recorded only four times from Western
Australia; once from the Williams River, where it was originally
obtained by Gilbert, and three times since from around Kojonup."
Glauert (1933, p. 19) gives its range in Western Australia as "Lower
South-West from Narrogin to Kojonup"; he adds that it "seems to
be rather rare, six specimens only having reached the [Perth]
Museum within the last five years."
For Central Australia Spencer (1896, p. 30) records only a single
specimen, taken at Alice Springs, and remarks that it "is evidently
not a common form in the' central district."
"The measurements given in the British Museum Catalogue of
1888 are taken from an Adelaide specimen, but I have failed to
trace any recent records of the animal in South Australia. . . .
To-day it is impossible to define its former range in the State, or,
unfortunately, even to attest to its present existence." (Jones,
1923, p. 102.)
Gould's statement of the range (1863, vol. 1, p. xxvii) as the
"interior of New South Wales and the colony of Victoria" is ob-
viously incomplete and supported by rather meager evidence. How-
ORDER MARSUPIALIA: MARSUPIALS 29
ever, Krefft states (1871 [p. 40]) that the animal occurs in New
South Wales, near the Darling River, and Iredale and Troughton
(1934, p. 8) include Victoria in the range.
On Williams River, Gilbert (in Gould, 1863, vol. 1, p. 39) records
the species as invading a storeroom. The type specimen was cap-
tured in that locality by a Domestic Cat.
"Some species, notably the brush-tailed and the lesser brush-
tailed phascogales, are now rather scarce over the greater part of
their range, having been greatly reduced by disease, which swept
off large numbers of native animals in 1898-9-1900. Cats have also
been very destructive." (Le Souef and Burrell, 1926, p. 333.)
Here we seem to have yet one more melancholy case of the virtual
disappearance of a species before any adequate knowledge of its life
history or even of its distribution was obtained.
Slender-tailed Pouched Mouse; Gray Pouched Mouse;
"Common" Pouched Mouse; Slender
Mouse-Sminthopsis
SMINTHOPSIS MURINA (Waterhouse)
This animal seems to have disappeared over considerable por-
tions of its original wide range in Australia. It has been divided
into the following four subspecies:
SMINTHOPSIS MURINA MURINA (Waterhouse)
Phascogale murina Waterhouse, Proc. Zool. Soc. London 1837, p. 76, 1838.
("North of Hunter's River, New South Wales.")
FIGS.: Waterhouse, 1841, pi. 10; Gould, 1863, vol. 1, pi. 43; Lydekker, 1894,
pi. 29 (subsp.?) ; Le Souef and Burrell, 1926, fig. 94.
Fur short and soft ; general color above gray, with a faint yellow-
ish tint; feet, under parts, and face beneath eyes white; tail covered
with minute silvery-white hairs. Head and body, 76 mm.; tail,
65 mm. (Waterhouse, 1838, p. 76.)
This form occurs in New South Wales and southern Queensland.
Waterhouse reported it from north of Hunter's River, New South
Wales, and Gilbert found it on the Severn River in the same state
(Gould, 1863, vol. 1, p. 50). It was perhaps this form that Gould
recorded (1863, vol. 1, p. 49 as Antechinus albipes) from the
Darling Downs of New South Wales.
This little marsupial, if sufficiently abundant, would evidently
act as a check on one of the introduced rodent pests. "Mr. A. C. V.
Bligh, of Toowoomba, Queensland, reports S. murina as being
numerous at the same time as the common mouse (M. musculus) ,
and feeding upon the latter" (Le Souef and Burrell, 1926, p. 355) .
30 EXTINCT AND VANISHING MAMMALS
SMINTHOPSIS MURINA ALBIPES (Waterhouse)
Phascogale albipes Waterhouse, Proc. Zool. Soc. London 1842, p. 48, 1842.
("Port Adelaide," South Australia.)
Fia.: Gould, 1852, vol. 1, pi. 42.
Upper parts brownish (the hairs being annulated with yellow
near the tip and with black at the tip) ; hairs of under parts deep
gray, tipped with white; feet white; tail dark, with very minute
hairs. Head and body, 95 mm.; tail, 80 mm. (Waterhouse, 1842,
p. 48.)
This subspecies occurs in South Australia and Victoria.
Gould gives it too wide a range in stating (1863, vol. 1, p. 49)
that it "appears to be almost universally distributed over the whole
of the southern coast of Australia, from Swan River to New South
Wales."
"These little animals . . . are caught in large numbers by the
aborigines of the Murray" (Krefft, 1871, p. [41]).
"Although in books this little animal passes uniformly under the
name of 'Common' Pouched Mouse, it is by no means a common
species. In South Australia it is not nearly so frequently met with
as is S. crassicaudata. ... It is an animal which is very rarely
seen except when it has fallen victim to a cat, and but little is
known of its life history." (Jones, 1923, p. 118.)
C. W. Brazenor (in litt., March 3, 1937) knows of no locality
in Victoria in which this animal can be found at the present time.
SMINTHOPSIS MURINA FULIGINOSA (Gould)
Antechinus juliginosus Gould, Mamm. Australia, vol. 1, pi. 41, 1852. ("At
King George's Sound and in the vicinity of Perth," Western Australia.
Thomas (1888, p. 305) lists the type specimen from "R. Avon, W. A.,"
which is in the general vicinity of Perth. Thus Iredale and Troughton
(1934, p. 10) are evidently in error in giving, as the restricted type locality,
"King George's Sound.")
FIG.: Gould, 1852, pi. 41.
Upper parts dark grayish brown, interspersed with longer black
hairs; face lighter; a mark around the eyes black; chest sooty gray,
with a narrow median line of buffy gray; rest of under parts pale
grayish white; feet buffy white; tail dark reddish brown, grayish
beneath. Head and body, 83 mm.; tail 83 mm. (Gould, 1863, vol. 1,
p. 48.)
The range is "South-Western Australia, more or less coastal, but
inland to Katanning, Broomehill, Gnowangerup, and Bulong, near
Kalgoorlie" (Glauert, 1933, p. 20) .
Gould (1863, vol. 1, p. 48) considered it "very abundant, both
at King George's Sound and in the vicinity of Perth."
ORDER MARSUPIALIA: MARSUPIALS 31
"These little animals ... are caught in large numbers by the
aborigines ... of King George's Sound" (Krefft, 1871, p. [41]).
Shortridge (1910, pp. 842-844; map, p. 843) speaks of it as
"occurring throughout the South -West; appears to be more plenti-
ful in the coastal districts wherever grass-trees (Xanthorrhoea)
occur ....
"On account of their habit of hiding among fallen timber or
tree-stumps, the marsupial mice must invariably get exterminated
wherever bush fires occur. This species, as well as Dromicia and
the small Phascogales, has consequently become very scarce, espe-
cially in the agricultural and more thickly populated areas. In addi-
tion it is probably to a great extent killed off by the cats that have
run wild in large numbers."
More recently, however, Glauert (1933, p. 20) considers it still
"a very common species in the South- West."
Gilbert (in Gould, 1863, vol. 1, p. 48) found it insectivorous.
SMINTHOPSIS MURINA CONSTRICTA Spencer
Sminthopsis murina var. constricta Spencer, Kept. Horn Sci. Exped. Central
Australia, pt. 2, zool., p. 33, 1896. ("Oodnadatta," South Australia.)
General coloration similar to that of S. m. murina; foot broader;
a small tuft of white hairs on posterior face of forearm; tail in-
crassated. Head and body, 71 mm.; tail, 80 mm. (Spencer, 1896,
p. 33.)
In his original description of constricta, Spencer mentions only
the single specimen from Oodnadatta, but on a previous page
(1896, p. 32) he records a specimen of "S. murina" from Alice
Springs, Central Australia, which perhaps belongs to the same
form. He adds that the species "does not appear to be common
in the central district."
No additional information concerning the present subspecies
seems to have come to light since its discovery was announced more
than 45 years ago.
Long-tailed Sminthopsis
SMINTHOPSIS LONGICAUDATA Spencer
Sminthopsis longicaudatus Spencer, Proc. Royal Soc. Victoria, n. s., vol. 21,
pt. 2, p. 449, 1909. ("West Australia.")
This little animal is known from only a single specimen, from
no more definite locality than "West Australia."
General body color gray, tinged with rufous in parts; a darkish
line through the eye; lips, chin, and feet white; tail scaly, with
32 EXTINCT AND VANISHING MAMMALS
short stiff hairs. Head and body, 100 mm.; tail, 202 mm. (Spencer,
1909, pp. 449-450.)
"This species does not seem to have been recorded since its
original discovery, and we have no information about the type
locality beyond Spencer's vague 'West Australia' " (Glauert, 1933,
p. 21).
The apparently total lack of additional information concerning
the species, during 30 years past, does not augur well for its present
status, although there is always a possibility that it may have
survived in some out-of-the-way corner of Western Australia.
Common Eastern Native Cat; Viverrine Native Cat
DASYURUS VIVERRINUS (Shaw)
Didelphis Viverrina Shaw, Gen. Zool., vol. 1, pt. 2, p. 491, pi. Ill, 1800. (A
composite species, based in part upon "The Tapoa Tafa" of White (Jour.
Voy. New South Wales, p. 281, pi. 58, 1790) and in part upon "The
Spotted Opossum" of Phillip (Voy. Botany Bay, p. 147, pi. 15, 1789).
The name has become restricted to the latter; type locality, Botany Bay,
New South Wales. Cf. Harper, 1940, p. 191.)
FIGS.: Waterhouse, 1841, pi. 7 (as D. maugei) ; Gould, 1863, vol. 1, pi. 50; Krefft,
1871, pi. 13; Lydekker, 1894, pi. 26; Le Souef and Burrell, 1926, figs. 87, 88;
Fleay, 1932, pis. 3, 4; Pocock, 1937, p. 616, fig.
This species, like its larger relative, Dasyurus maculatus, was
distributed through eastern Australia and Tasmania and has suf-
fered a similar or perhaps even greater reduction in numbers.
There are two color phases, of which the black is the less com-
mon. Fur thick and soft; general color either pale olive-gray or
deep black, profusely spotted with white; belly and limbs paler
than back; tail bushy, without spots, tipped with white in the gray
phase. Head and body, 400-440 mm.; tail, 210-290 mm. (Thomas,
1888, pp. 266-267.)
The range includes New South Wales, Victoria, South Australia,
and Tasmania.
The history and status of the species are reviewed by Jones (1923,
pp. 91-92) :
It was abundant round, and even in the immediate precincts of, the larger
Australian towns. Twenty years ago it was exceedingly common about Ade-
laide. Still more recently it lived close to Melbourne; and to-day it is not
uncommon in the suburbs of Sydney. . . . Very early in the days of colonisa-
tion it was regarded with dislike because of the damage it did by killing
poultry; but there are many settlers who would now welcome its return in
order to keep the mice plagues within check. . . .
There is no doubt that as a destroyer of mice, rats, and young rabbits the
Native Cat played an extremely useful part in Australian rural economy,
and despite the fact that it was an occasional robber of hen roosts its presence
was a real asset to the country.
ORDER MARSUPIALIA I MARSUPIALS
33
Its range in South Australia was formerly very wide. On Kangaroo Island
it appears to have been always more or less of a rarity. Thirty years ago it
haunted the shores of the [Murray] river and lakes, being there very partial
to a fish diet. To-day, if it exists at all in this State [South Australia], it
must be an animal of the utmost rarity. Although there is no doubt that
the influences which have been at work in the general process of the extermina-
tion of the Australian fauna have operated to the full on the Native Cat;
it is possible that another factor has come into play during the final scene
of its passing. The animal has been trapped, poisoned and persecuted through-
out the country .... The Native Cat, with its cunning and its activity, was
FIG. 1. Common Eastern Native Cat (Dasyurus viverrinus)
well able to look after itself, despite the fact that it was an extremely easy
animal to trap. Its rapid decrease started about the year 1900, and during
that and the two following years the so-called "common" Native Cat practically
disappeared from South Australia. Much the same thing happened in Victoria
and in New South Wales, with the exception of the district immediately
round Sydney. It would seem that some epidemic disease must have spread
through the Dasyures, and that after a lapse of twenty years the remnant
has not succeeded in re-establishing itself. In the Animal Protection Act of
1919 the Native Cat is not even mentioned. The evil or the good that it did
has ceased to be a factor of any economic importance.
At the present time it "still haunts the coastal cliffs and moun-
tains about Sydney, also parts of Tasmania, Victoria, and New
South Wales, where survival seems assured" (E. Le G. Troughton,
in Hit., April 16, 1937) .
In Victoria, according to C. W. Brazenor (in litt., March 3, 1937),
it occurs in greatest numbers around Lake Corangamite but is also
34 EXTINCT AND VANISHING MAMMALS
seen occasionally in the eastern part of the state. Further informa-
tion is supplied by Fleay (1932, pp. 63-66) :
From accounts supplied by people of these localities [about Lake Coranga-
mite], the animals were not affected by the mysterious disease which an-
nihilated many marsupials in other parts of the country in the first years of
this century. Though still well known, however, they are becoming scarce,
with the continuous work of rabbiters' dogs and traps, and the increase in
settlement. . . .
The adult males supported a host of parasites, and, when first brought to
Melbourne, before being treated, they possessed numerous large ticks, sores
infested with fly larvae, and the peculiar flea (Stephanocercus dasyuri) ;
while investigations on Dasyures which had been caught in rabbit traps
showed numerous nematode worms internally.
On Kangaroo Island, South Australia, the species seems to have
disappeared (Waite and Jones, 1927, p. 322) .
In Tasmania it has fared better than in Australia. "The common
Dasyure ... is to be met with in many localities in spite of the
warfare waged against them in return for the toll they take of the
settlers' poultry. In this respect, however, they cannot be con-
sidered so destructive as the Tiger Cat' [D. maculatus]" (Lord
and Scott, 1924, p. 270.) "The Dasyure is scattered throughout
Tasmania, and still is very plentiful. This marsupial does not seem
to either increase or decrease." (R. Boswell, in litt., May 13, 1937.)
Despite Jones's statement (1923, p. 92) that "the animal's skin is
of no commercial value," there evidently was a demand for it in
former years. Lydekker writes (1894, p. 164) : "The fur being soft,
the skins are suitable for linings; and from two to five thousand
skins are annually imported into England. Formerly the grey skins
fetched from about fivepence to sixpence each in the market, while
the value of the black ones ranged from tenpence to a shilling. Of
late years, however, there has been a fall in the price."
The species is now under complete legal protection in Victoria
and in Tasmania.
Geoffrey's Native Cat; Black-tail CM! Native Cat
DASYURUS GBOFFROII GEOFFROII Gould
Dasyurus Geoffroii Gould, Proc. Zool. Soc. London 1840, p. 151, 1841. ("Liver-
pool Plains/' New South Wales.)
FIGS.: Gould, 1851, vol. 1, pi. 51; Cabrera, 1919, pi. 4, fig. 2.
The typical subspecies of Geoffrey's Native Cat seems to have
become extinct over the greater part of its range, but it may sur-
vive in parts of New South Wales and Queensland.
General color fuscous, washed with yellow; head, back, and sides
with white spots, smaller than those in D. viverrinus and D. macu-
ORDER MARSUPIALIA: MARSUPIALS 35
latus; tail long, terminal half black; under parts whitish. Head
and body, 15 inches; tail, 11.5 inches. (Gould, 1841a, p. 151.)
The former range of this animal extended from Victoria and
South Australia through New South Wales to Queensland but
apparently did not include the coast region of the southeast or the
extreme north.
According to Gould (1863, vol. 1, p. 58), the species (including
both subspecies) "inhabits the whole of the southern portion of the
country from Moreton Bay [Queensland] on the east to Swan
River on the west." It "appears to be exclusively confined to the
regions on the interior side of the hills, the specimens I have seen
having been procured on the Liverpool Plains in New South Wales,
the Murray Scrub in South Australia, and beyond the ranges of
Swan River on the western coast."
In Victoria it was always confined to the northwestern corner.
The last known record was in 1857, and the species is now extinct
in that state. (C. W. Brazenor, in Hit., March 3, 1937.)
In South Australia there is no record other than that of Gould
and a specimen listed in the British Museum Catalogue of 1888.
"Men who have been professionally interested in the fauna of the
State for a period of forty years are unaware of any examples being
taken in South Australia proper. Unless it still lingers near to the
northern limits of the State, it must probably be regarded as extinct
in South Australia." (Jones, 1923, pp. 93-94.)
In the Dawson Valley of Queensland, in 1905, it "was noticed to
be suddenly numerous, but it completely vanished by 1906" (Fin-
layson, 1934, p. 225) . It is represented in the Queensland Museum
merely by two specimens without precise localities (Longman, 1930,
p. 62).
The eastern subspecies has very likely suffered in the same way
as the western, which was "killed off as much as possible in the
agricultural and more thickly populated districts on account of
being so destructive to poultry" (Shortridge, 1910, pp. 838-839).
Hoy (1923, p. 165) contributes information on an important enemy:
"I . . . am told that domestic cats frequently kill and drag home
adult native cats (Dasyurus viverrinus, D. geoffroyi, and D. hallu-
catus)" Other possible causes underlying the sudden fluctuations
in numbers of the species and its general disappearance over most
of its range, have not been definitely explained.
[The larger western subspecies (D. g. fortis Thomas) still occurs
commonly in the southwest of Western Australia (Shortridge, 1910,
pp. 837-839; Glauert, 1933, p. 18, and in litt., March 17, 1937).
Some representative of the species perhaps fortis is reported from
Central Australia but as nowhere common there (Finlayson, 19356,
36 EXTINCT AND VANISHING MAMMALS
pp. 60-61). If this is the western subspecies, the eastern part of
Western Australia forms a great blank in its known distribution.]
Large Spotted-tailed Tiger-cat; Spotted-tailed Dasyure
DASYURUS MACULATUS (Kerr)
Viverra maculata Kerr, Anim. Kingdom of Linnaeus, p. 170, 1792. (Based
upon the "Spotted Martin" of Phillip, Voy. Botany Bay, p. 276, pi. 46,
1789; type locality "the neighborhood of Port Jackson" [Sydney], New
South Wales.)
FIGS.: Waterhouse, 1841, pi. 6 (as D. macrowrus} ; Gould, 1851, vol. 1, pi. 49;
Lydekker, 1894, pi. 25; Raven, 1924, p. 25; Le Souef and Burrell, 1926,
fig. 86; Fleay, 1932, p. 66, fig. 4, and pi. 5.
This fierce and rather powerful animal, one of the largest of the
carnivorous marsupials, is found in eastern Australia and in Tas-
mania. Its range and its numbers have been reduced by settlement,
though evidently not yet to the danger point.
According to Phillip (1789, p. 276), the general color is black;
body and tail irregularly blotched with white; tail tapering to a
point; head and body, 18 inches; tail, nearly 18 inches. But Water-
house (1846, pp. 440-441) and later authorities do not agree with
Phillip and Kerr on the general color; it "varies from a very deep
brown to a rich red-brown"; under parts "dirty yellow"; head and
body, 17-24 inches; tail, 15-20 inches.
The range includes "south-eastern Queensland, eastern New South
Wales, Victoria, south-eastern South Australia, Tasmania" (Iredale
and Troughton, 1934, p. 14) . Some of the earlier works extend the
range into central or northern Queensland. According to Le Souef
and Burrell (1926, p. 322), the species "is fairly common in Eastern
Australia, from Cape York to Victoria." Half a century ago Thomas
(1888, p. 265) considered it "approaching . . . complete extermina-
tion in Australia"; but Ogilby (1892, p. 18) replied that it "is by
no means uncommon nor seemingly has it any present intention
of dying out in the mountainous and coastal districts of eastern
Australia." On the Comboyne Plateau of New South Wales "it
appears to be rather uncommon" (Chisholm, 1925, p. 72) .
In Victoria it was "common in heavily scrubbed country till
about 1907, at which time an epidemic of disease almost com-
pletely destroyed the species. Has recovered somewhat in recent
years and is found in some numbers in the Otway Ranges, and to a
lesser extent scattered throughout the Dividing Range." (C. W.
Brazenor, in litt., March 3, 1937.) "With the advent of settlement,
disease, dogs, guns, traps, and . . . the fox, which exterminates
the simple marsupial game of the Dasyure, we have come to the
time, in Victoria, of the almost complete disappearance of these
ORDER MARSUPIALIA: MARSUPIALS 37
primitive carnivorous hunters" (Fleay, 1932, p. 68). The species
now has complete legal protection in Victoria.
"Probably never abundant in South Australia, the stronghold of
the species was in the south-eastern portion of the State. It is
possible that some few still exist in the less closely settled areas
of the South-East." (Jones, 1923, p. 88.)
In Tasmania it "is regarded as one of the settlers' greatest pests,
owing to the toll it will take of his poultry" (Lord and Scott, 1924,
p. 269). "The enemy of the settler's chickens, it is only natural
that this species should be reduced in numbers, especially in the
settled districts. Even so, this hardly accounts for the scarcity of
this species in the more Southern Tasmanian localities in the last
few years. In the North-West the species is still fairly common."
(Lord, 1928, p. 22.)
There are additional records of nocturnal raids on poultry on
the mainland of Australia, and this habit naturally reacts against
the species. "All three dasyures are doomed to extinction, since
they are killed whenever met with by the man on the land" (Jones
and Manson, 1935, p. 34) . "It is now being replaced by the domestic
cat and the fox" (Raven, 1924, p. 25) . However, "it is able to kill
wallabies and fairly large birds," and "one succeeded, after a
severe battle, in killing a large tom-cat" (Le Souef and Burrell,
1926, pp. 322-323). Although Lydekker wrote in 1894 (p. 160) that
"its skin is but little valued by furriers," it must be remembered
that many furs, formerly in little demand, now bring good prices.
While the Dingo is generally considered responsible for the ex-
tinction of the terrestrial Tasmanian Devil and Tasmanian Wolf in
Australia, opinion seems divided as to whether it has seriously
affected the status of the arboreal Spotted-tailed Tiger-cat on the
mainland. E. Le G. Troughton (in litt., April 16, 1937) believes that
this species, by reason of its furtive and aggressive disposition,
should survive indefinitely in the dividing ranges of the east coast.
Slender Native Cat; Slender Spotted-tailed Tiger-cat
DASYURUS GRACILIS Ramsay
Dasyurus gracilis Ramsay, Proc. Linn. Soc. New South Wales, ser. 2, vol. 3,
p. 1296, 1888. ("Bellenden-Ker Ranges," northern Queensland.)
The Slender Native Cat is one of those species which, as far as
known, has always been very rare. Apparently less than half a
dozen specimens are on record all from northern Queensland.
General color, above and below, deep blackish brown with white
spots; tail spotted and closely furred, with a terminal tuft on the
upper side. Total length, about 23 inches; tail, 9.3 inches. (Ramsay,
op. tit., p. 1296.)
38 EXTINCT AND VANISHING MAMMALS
The type specimen was collected by Robert Grant on the Bel-
lenden-Ker Ranges, apparently in 1887. The species was next
found years later by C. M. Hoy on the Atherton Tableland (Le
Souef and Burrell, 1926, p. 324) . The late Henry C. Raven, of the
American Museum of Natural History, informed me that he secured
two or three specimens about 1922 in the same general region.
Whatever the factors may be that seem to restrict so decidedly
the numbers of the Slender Native Cat, they have not been ascer-
tained.
According to E. Le G. Troughton (in litt., April 16, 1937) , the
species is rarely captured because of the density of its mountain
rain-forest habitat, and should therefore survive in parts of coastal
Queensland for all time.
Tasmanian Devil
SARCOPHILUS HARRISII (Boitard)
Didelphis ursina Harris, Trans. Linn. Soc. London, vol. 9, p. 176, pi. 19,
fig. 2, 1808. ("Van Diemen's Land.") (Not Didelphis ursina Shaw (1800).)
Ursinus harrisii Boitard, Jardin des Plantes, p. 290, "1842" = 1841. (Tasmania.)
FIGS.: Geoffroy and Cuvier, Hist. Nat. Mammif., vol. 7, pi. 113, 1842;
Gould, 1851, vol. 1, pi. 48; Royal Nat. Hist., vol. 3, p. 271, fig., 1894-95;
G. Smith, 1909, fig. 24; Raven, 1924, p. 25, fig., and 1929, p. 204, fig.;
Le Souef and Burrell, 1926, fig. 85; Fleay, 1935, pi. 9; Pocock, 1937,
p. 615, fig.; Reed and Lucas, 1937, p. 89, fig. 33.
This fierce little beast occurred in past ages on the Australian
mainland, where it presumably succumbed to the advancing Dingo.
In 1912 a specimen, probably an escaped captive, was taken in
Victoria. It "is now confined to Tasmania, where it maintains a
rather precarious foothold in the wilder parts of the country"
(Jones, 1923, p. 85).
Whole body and upper part of tail covered with long coarse
black hair; irregular blotches of white on shoulders, throat, or
rump (G. P. Harris, 1808, p. 176) . It is a thickset, powerful animal,
and, except for its tail, resembles a miniature bear in outline. Head
and body, 670-825 mm.; tail, 258-300 mm. (Lord and Scott, 1924,
p. 267.)
"These animals were very common on our first settling at Hobart
Town, and were particularly destructive to poultry, &c. They,
however, furnished the convicts with a fresh meal, and the taste
was said to be not unlike veal. As the settlement increased, and the
ground became cleared, they were driven from their haunts near
the town to the deeper recesses of the forests yet unexplored." (G. P.
Harris, 1808, p. 177.)
"The devil is destructive to sheep all over the colony, and is
ORDER MARSUPIALIA: MARSUPIALS
39
indeed the most destructive of our indigenous quadrupeds, the
Thylacinus being much scarcer" (Gunn, 1838, p. 104).
"It has now become so scarce in all the cultivated districts, that
it is rarely, if ever, seen there in a state of nature; there are yet,
however, large districts in Van Diemen's Land untrodden by man;
and such localities, particularly the rocky gullies and vast forests
on the western side of the island, afford it a secure retreat. . . .
"In its disposition it is untameable and savage in the extreme,
and is not only destructive to the smaller kangaroos and other
native quadrupeds, but assails the sheep-folds and hen-roosts when-
ever an opportunity occurs." (Gould, 1863, vol. 1, p. 55.)
FIG. 2. Tasmanian Devil (Sarcophilus harrisii)
"The Devil is far commoner than the Tiger and more widely dis-
tributed through the island .... Like the Tiger it destroys sheep,
making a single meal off each capture." (G. Smith, 1909, p. 97.)
Lord (1928, p. 22) says of it:
The Tasmanian Devil will probably survive for many years. Its hardy
nature both in captivity and in its wild state cause [s] one to wonder how it
came about that this species became extinct on the mainland within com-
paratively recent geological times. It cannot be considered a pleasant animal
to have much to do with, and numbers are killed by trappers in the course
of their work. In the rougher sections of the country this species exists in
fair numbers and there is every prospect of it remaining an inhabitant of
such places for years to come.
One or more Tasmanian Devils will often follow a Thylacine on its hunting
excursions. The Thylacine will kill a wallaby or other small animal, select a
few choice morsels, and pass on. The Devils will carry on the feast and
consume the remnants, bones and all.
According to R. Boswell (in Hit., May 13, 1937), it still exists
in large enough numbers to enable it to be out of immediate danger
40 EXTINCT AND VANISHING MAMMALS
of extinction. Civilization has been the great cause of its decreasing
numbers. It has no legal protection.
"In spite of his ungainly, ugly appearance, his whining snarls
and unpleasant smell, the Tasmanian Devil is a creature of many
amusing antics and distinctly unusual ways. Moreover, his position
as the second largest of living marsupial carnivores, soon, perhaps,
to be the largest when the rare Thylacine finally disappears, invests
him with a peculiar interest." (Fleay, 1935, p. 100.)
Tasmanian Wolf; Marsupial Wolf; Tasmanian Tiger;
Thylacine
THYLACINUS CYNOCEPHALUS (Harris)
Didelphis cynocephala Harris, Trans. Linn. Soc. London, vol. 9, p. 174, pi. 19,
fig. 1, 1808. ("Van Diemen's Land" [= Tasmania].)
FIGS.: Waterhouse, 1841, pi. 5; Gould, 18*51, vol. 1, pis. 53, 54; Wolf, 1861,
pi. 31; Krefft, 1871, pi. 12; Royal Nat. Hist., vol. 3, p. 270, fig., 1894-95;
G. Smith, 1909, fig. 23; Cabrera, 1919, pi. 6; Australian Mus. Mag., vol. 1,
no. 3, p. 62, frontisp., 1921; Le Souef and Burrell, 1926, fig. 84; Raven,
1929, p. 207, fig.; Pocock, 1937, p. 614, fig.; Reed and Lucas, 1937, p. 85,
fig. 31; Sharland, 1939, p. 23, fig.
This largest and most formidable of living carnivorous marsupials
is so seriously reduced in numbers that its fate seems to be hanging
by a somewhat slender thread.
General build doglike, but hind end tapering gradually to the
tail; upper parts tawny grayish brown, with 16-19 blackish brown
bands across the back, chiefly developed on the hind quarters;
under parts paler. Head and body, 1230-1300 mm.; tail, 525-650
mm.; height at shoulders, about 560 mm. (Chiefly from Lord and
Scott, 1924, p. 264.)
While a fossil form of Thylacine has been recorded from the
Australian mainland, the range of the living form is restricted to
Tasmania. The mainland Thylacine is presumed to have suc-
cumbed as a consequence of the advent of the Dingo during the
Pleistocene, for it probably could not compete successfully with
that more highly organized animal.
The Thylacine "is common in the more remote parts of the
colony, and they are accordingly often caught at Woolworth and
the Hampshire hills. . . . They are usually nocturnal in their
attacks on sheep." (Gunn, 1838, p. 101.)
It was with prophetic vision that Gould wrote long ago (1863,
vol. 1, pp. 60-61) :
When the comparatively small island of Tasmania becomes more densely
populated, and its primitive forests are intersected with roads from the eastern
to the western coast, the numbers of this singular animal will speedily diminish,
ORDER MARSUPIALIA : MARSUPIALS 41
extermination will have its full sway, and it will then, like the Wolf in
England and Scotland, be recorded as an animal of the past: although this
will be a source of much regret, neither the shepherd nor the farmer can be
blamed for wishing to rid the island of so troublesome a creature. A price
is already put upon the head of the native Tiger, as it is called; but the
fastnesses of the Tasmanian rocky gullies, clothed with impenetrable forests,
will, for the present, preserve it from destruction.
. . . Although too feeble to make a successful attack on man, it commits
sad havoc among the smaller quadrupeds of the country, and among the
poultry, and other domestic animals of the settler; even sheep are not secure
from its attacks ....
"The damage which it inflicts on the flocks of the settlers has
. . . given rise to a relentless war of extermination, which has
resulted in the almost complete extinction of this, the largest of
the Australasian Carnivores, in the more settled portions of the
country" (Lydekker, 1894, p. 152).
G. Smith (1909, pp. 96-97) wrote:
The destructiveness of these animals is greatly enhanced by the fact that a
Tiger will make only one meal of a sheep, merely sucking the blood from
the jugular vein or perhaps devouring the fat round the kidneys, but it
never returns to the same carcass. . . . The shepherds wage incessant war
on the creature, in the summer laying traps and hunting it with dogs, in the
winter following up its tracks through the snow. A reward of a ' pound is
given for the head by the Government, but the shepherd generally rides
round with the head to several sheep-owners in the district, and takes toll
from them all before depositing it at the police station. In consequence a
large reward must be offered for the carcass of a Tiger, and an offer of 10
during a year for a live Tiger to be delivered in Launceston was unsuccessful.
It pays the shepherd very much better just to hack off the head and take it
round on his rides. Although the Tiger is by no means confined to the Lake
District, it is more abundant here than anywhere else, though a stray individual
may turn up on nearly all the big sheep stations throughout the island.
Lord (1928, pp. 20-21) says of the Thylacine:
The animal is confined practically to the rugged western portion of the
island. From the more settled districts it has long since disappeared, and
even in the more distant sheep runs it has been trapped out .... It is
now also being killed out even in the rugged and more inaccessible parts
of the country, which tends to reduce still further the remnants of this
species. The explanation of this is that the Thylacine interferes with the
trappers' snares. As a result, a powerful "springer" snare is set often in the
vicinity of their "skinning yards," which are situated every quarter of a
mile or so along the lines of snares. Thylacines or other animals caught
in these powerful snares are, as a rule, too severely injured to be kept alive as
specimens for zoological gardens, even if the trappers would take the trouble
to bring them in. The extended trapping of recent years will tend, therefore,
to restrict the Thylacine to the most rugged and unsettled portions of the
West of the island. Here it may survive as a living species for years to come,
but its eventual doom seems apparent unless such attempts as are being made
at present by Mr. A. R. Reid (Curator of the Beaumaris Zoo, Hobart) to
breed these animals in captivity are successful. . . .
It is doubtful if the shy animal will breed within the confines of a Zoo,
and it would be in the interests of science if a reserve could be set aside and
42
EXTINCT AND VANISHING MAMMALS
netted in in order to prevent total extermination. ... If funds were available
an area in the National Park might well be considered for such a reserve.
"The Tasmanian Tiger is now only to be met with in a very few
numbers. This animal is causing great concern in Tasmania at
the present time. It is thought by many to be extinct, but this is
not so. I have obtained authentic reports regarding its presence
as having been seen as recently as January 1937 on the West
Coast of Tasmania.
\
FIG. 3. Tasmanian Wolf (Thylacinus cynocephalus)
11 The former range of the Tasmanian Tiger must have been very
great as I know of one Tasmanian, who with his brother, killed as
many as twenty-four of these animals during one day, and received
a reward of 1. 0. per head for each animal.
"The Tasmanian Tiger is now wholly Protected." (R. Boswell,
in litt., May 13, 1937.)
"The significance of the mainland elimination, prior to settlement,
of the largest living marsupial carnivore (Thylacinus) has already
been noted, and latest reports from Tasmanian authorities indicate
grave doubts for the insular survival of this unique example of
parallelism" (Troughton, 1938, p. 408) .
However, the latest news from Tasmania is distinctly encouraging.
It comes in the form of a paper by Sharland (1939), which gives
an account of several recent expeditions that have been sent to
mountainous areas in the western part of the state by the Tasmanian
Animals and Birds' Protection Board. From this account the fol-
lowing information is derived:
"The Thylacine exists to-day as but a remnant of the numbers
ORDER MARSUPIALIA: MARSUPIALS 43
which, 50 or 60 years ago, roamed the countryside, feeding on small
marsupials and sheep. . . . Nowadays, certainly, it is rarely seen.
. . . When the game season is opened every few years the animal is
often caught in snares. But it is in no part specially common, and
there are extensive areas in this region where it does not occur at
all, or but sparsely, its distribution depending almost wholly on the
presence of smaller 'game.' " (P. 20.)
"The Thylacine has been known to attack dogs when cornered,
but so far as I can determine there is no record of its ever having
attacked man" (p. 32) .
In a great amphitheatre about 25 to 40 miles in diameter, bounded
by the King William, Prince of Wales, Norway, and other ranges,
"we came upon many tracks made by Thylacine, indicating that the
animal was fairly common and well distributed" (p. 34) .
"The area enclosed by the mountains would make a splendid game
sanctuary .... The Thylacine is probably as common here as in
any other part of the West Coast." (P. 34.)
Additional tracks were found in the Jane River region, where
"the animal had apparently been trailing Wallaby" (p. 34).
"No longer a menace to sheep-owners since its isolation in the
remote parts of the State, the animal possesses a unique scientific
value which is appreciated by the Board. While, up to half a
century ago, it was fairly plentiful in the grazing country of the
central plateau, and was known also to inhabit parts of the eastern
tiers and other mountain forest areas adjacent to settlement, it has
now practically disappeared from these districts, to make its last
stand in the western section of the State." (P. 36.) The recent
opening of a road through this remote region has had an adverse
effect upon the Thylacine's prospects for survival.
A mountainous area situated about Frenchman's Gap, east of
Macquarie Harbor, and comprising approximately 300,000 acres,
is suggested as a suitable sanctuary for the Thylacine and other
animals (p. 38) .
Family MYRMECOBIIDAE : Marsupial Anteaters
The single genus of this family consists of two forms, both of
which are treated here. They occur in the southern half of Australia.
Banded Anteater; West Australian Numbat
MYRMECOBIUS PASCIATUS FASCIATUS Water-house
Myrmecobius fasciatus Waterhouse, Proc. Zool. Soc. London 1836, p. 69,
1836. ("In the interior of the Swan River Settlement, about 90 miles to
the S.E. of the mouth of that river," Western Australia.)
44 EXTINCT AND VANISHING MAMMALS
FIGS.: Waterhouse, 1838a, pi. 27, and 1841, pi. 11; Gould, 1845, vol. 1, pi. 4;
Lydekker, 1894, pi. 30; Royal Nat. Hist., vol. 3, p. 275, lower fig., 1894-95;
Cabrera, 1919, pi. 7; Le Souef and Burrell, 1926, fig. 96.
This beautiful little animal, representing a special family related
to the dasyures, has been undergoing shrinkage of range and reduc-
tion in numbers in Western Australia for a considerable period and
is perhaps approaching extinction.
Color above reddish ochre, interspersed with white hairs ; posterior
half of body with alternate black and white bands; tail long-haired,
mixed with black, white, and reddish ochre; legs chiefly pale buff;
under parts yellowish white (Waterhouse, 1836, pp. 69-70). Form
graceful, squirrellike ; a black stripe through the eye, and a white
stripe above it. Head and body, 220-240 mm.; tail, 160-175 mm.
(Thomas, 1888, pp. 311-314.)
In earlier days its range extended west to the Darling Range,
northwest to the vicinity of Moore's River, northeast to Laverton,
east to Kalgoorlie and possibly to South Australia near the coast,
and south to the vicinity of Albany (Shortridge, 1910, p. 846, map;
Glauert, 1933, p. 22). Forty years ago it was "fairly numerous
throughout the South-west, especially where the prevailing timbers
are the white gum (Eucalyptus redunca) and the jam (Acacia
acuminata) , getting less plentiful outside that area" (Shortridge,
in Thomas, 1907, p. 772) . "The Western Australian animal is now
excessively rare, and it is probable that before many years are
passed it will follow its South Australian neighbour [M. f. rujus]
into extinction" (Jones, 1923, p. 126). Troughton writes (1923,
p. 155) that "this animal . . . can now only be found in a greatly
restricted area"; he secured three specimens about 1921. It seems
to survive chiefly in the southwestern corner of the state, between
Perth and Albany.
"The beautiful little Banded ant-eater is much sought after on
account of its skin" (W. H. D. Le Souef, 1907, p. 406) .
Le Souef and Burrell (1926, pp. 365-366) write:
Quiet, inoffensive, without means of defence or offence, it is remarkable
that the marsupial ant-eater has survived through the ages. This could
happen only in Australia, where it did not come into competition with the
more advanced forms of life. . . .
It is abroad both by day and by night, and, being conspicuous and not at
all speedy, it makes a fairly easy mark for predacious animals and birds,
more especially the introduced cat and fox; to say nothing of the settlers'
dogs. Consequently, it is one of the first animals to disappear before the
inevitable opening up of the country, and it is now scarce over the greater
part of its range.
"The typical race ... is endangered by clearing, fires, and intro-
duced pests, the advent of the fox alone probably spelling the ulti-
ORDER MARSUPIALIA: MARSUPIALS 45
/
mate doom of the terrestrial and non-burrowing highly specialized
creature. . . . Myrmecobius may be regarded as one of the marsu-
pials within sight of extermination, in this instance not due to
exploitation by man, but as the result of settlement and introduced
enemies. Hope for ultimate survival may rest with the introduction
of a healthy colony to an island providing adequate supplies of
favoured diet, and absence of enemies." (E. Le G. Troughton,
in Hit., April 16, 1937.)
Its "very existence ... is threatened by both fox and cat"
(Troughton, 1938, p. 404) .
South Australian Numbat; Rusty Numbat
MYRMECOBIUS FASCIATUS RUFUS Jones
Myrmecobius rujus Jones, Mammals S. Australia, pt. 1, p. 123, figs. 79, 83, and
84, 1923. ("South Australia.")
FIG. : Jones, 1923, fig. 79.
This form of Numbat, apparently extinct in New South Wales
and coastal South Australia, still lingers in northwestern South
Australia and in southwestern Central Australia.
It differs from the West Australian Numbat in having the darker
part of the lower back "a fine bright brown" instead of blackish; tail
"a uniform grizzle of rust red and dark brown." Head and body,
175 mm.; tail, 135 mm. (Jones, 1923, pp. 124-126.) Finlayson
states (1933c, p. 204) that the outer surface of the ear is bright
rufous instead of yellow and black, and he gives the following
measurements for specimens from northwestern South Australia:
head and body, 200-270 mm.; tail, 130-170 mm.
"The New South Wales animal, reported fairly plentiful about
the plains of the Murray and Darling Rivers in 1862, ... is
apparently extinct" (E. Le G. Troughton, in litt., April 16, 1937).
It is perhaps the present form to which Helms refers (1896,
p. 255) in reporting the observations of the Elder Expedition
somewhere in South or Western Australia: "A more exciting piece
of work [by the natives] than digging for lizards is the excavating
for the quick, little, banded anteater, Myrmecobius jasciatus, which
animal often makes its lair over three feet below the surface."
This expedition brought back a dried skin from the Everard Range,
South Australia (Stirling and Zietz, 1893, p. 154) .
Jones (1923, pp. 126-127) says of this Numbat:
The Numbat was probably never a very abundant animal, but its distribu-
tion was comparatively wide. Only twenty years ago it was met with along
the scrub lands of the Murray, and earlier than that it existed quite near to
Adelaide. Enquiries as to its present existence have produced negative replies
from all those parts of the State in which there are schools, and the circulation
46 EXTINCT AND VANISHING MAMMALS
of its picture and description to more remote districts have proved equally
unavailing. The aboriginals who are attracted to civilisation, as it is represented
by the East-West railway, know the animal, but so far have failed to supply
any evidence as to its actual existence at the present time. If the Numbat
still exists in South Australia it is probably towards the Western Australian
border, and here it is probably the Western Australian form. The characteristic
South Australian type has probably gone for ever. . . . The extermination
of the Numbat is a tragedy in which man has probably played very little
conscious part; it is no tale of ruthless slaughter for gain, such as is being
rehearsed to-day in regard to the Australian fur-bearing animals, nor is it a
case of determined persecution as is the case with the Tasmanian Devil.
MyrmecobiiLS is an animal which is probably phylogenetically senile, which
FIG. 4. South Australian Numbat (Myrmecobius jasciatus rujus).
After Jones, 1923.
has become highly specialised in function and degenerate in some details
of structure. Added to this is the fact that its home is invariably made in
the hollow of a fallen tree or a rotting log. Accidental bush fires and the
intentional burning off of country seem to have found the Numbat an easy
victim, and they have exterminated it as they are exterminating other small
terrestrial Marsupials. There is no escape from a bush fire for the Numbat.
It does not excavate deep burrows, it does not climb, it is not fleet of foot
as its log home burns, it perishes. . . .
It is surely a tragedy that this most interesting animal has probably passed
out of existence in our State, and is rapidly repeating the process in a
neighbouring one without any representative collection having been made of
its remains. It will not be long before Myrmecobius will be as extinct as those
Mesozoic Marsupials of the English Jurassic beds of which it has been said
to be "actually an unmodified survivor."
Since the publication of Jones's account, investigation by Finlay-
son has shown that the species still survives in the arid center of
the continent. Rewrites (1933c, p. 203) :
"Its presence in the centre [in the Everard Range] was first
established by the work of the Elder Expedition ....
ORDER MARSUPIALIA: MARSUPIALS 47
"Recent field work ... in the far north-west of this State [South
Australia] (in a typical eremian environment) has shown . . .
that Myrmecobius still has a wide distribution in the south-west
parts of the centre beyond the limits of pastoral settlement, and in
some localities is by no means uncommon. It is possible that these
colonies actually link up with the far south-western ones in Western
Australia in a continuous band of distribution."
Finlayson here proposes (1933c, p. 203) to separate the central
animal from that of Western Australia under the name of Myrme-
cobius fasciatus var. rufus and gives (p. 204) as type locality
"mulga sand dunes, south and south-west of the Everard Range, far
north-west of State of South Australia." This name, however, is
antedated by Myrmecobius rufus Jones (1923), which was intro-
duced without any formal designation of type locality, but which
was based upon "South Australian specimens, from the Murray
and from near Adelaide" (Finlayson, 1933c, p. 205). The range,
according to Finlayson (p. 204), is "at present apparently not
north of about 25 S. lat., nor east of 132 30' E. long. To the south
and west as yet undetermined. Formerly as far south as Adelaide,
and probably ranging east into the Victorian and New South Wales
mallee areas."
Family PERAMELIDAE : Bandicoots
The range of the bandicoots extends over Australia, Tasmania,
New Guinea, and certain adjacent islands. There are about 9
genera, represented by about 44 forms. Of the latter, accounts of
12 appear in the following pages.
Eastern Barred Bandicoot ; New South Wales Barred Bandicoot
PERAMELES FASCIATA J. E. Gray
Perameles fasciata J. E. Gray, in Grey, Two Expeditions Australia, vol. 2,
appendix, pp. 401, 407, 1841. ("Liverpool Plains and South Australia";
type locality restricted by Thomas (1922, p. 144) to "Liverpool Plains,"
New South Wales.)
Fia: Gould, 1849, vol. 1, pi. 8.
This bandicoot occurred formerly in New South Wales and
Victoria. It has not been recorded for many years, however, and
is probably extinct (A. S. Le Souef, in litt., February 15, 1937).
This species has been more or less confused in descriptions with
P. myosura notina. "Grey brown, rump with three black bands;
tail white, with a black streak along the upper side. . . . Smaller
than P. Gunnii." (J. E. Gray, in Grey, 1841, p. 407.) Upper parts
penciled with black and yellow; sides yellow; under parts and feet
48
EXTINCT AND VANISHING MAMMALS
white (Gould, 1863, vol. 1, p. 12). Outer surface of ears flesh color
basally, darker terminally; sides of rump with four pale vertical
bands running downward from near the middle line, the spaces
between them brown or black (Thomas, 1888, p. 248) .
"This elegant species . . . enjoys a wide range over the eastern
. . . portions of Australia, but is more frequently met with in the
country within the ranges . . . than in the districts lying between
the mountains and the sea. In New South Wales, the stony ridges
which branch off from the ranges towards the rivers Darling and
Namoi, are localities in which it may always be found." (Gould,
1863, vol. 1, p. 12.)
IE. .
FIG. 5. Eastern Barred Bandicoot (Perameles jasciata). After Gould, 1849.
The animal is "now believed extinct though once well distributed
over western Victoria and N. S. Wales. The ultimate fate of these
small non-burrowing forms is most uncertain." (E. Le G. Trough-
ton, in litt., April 16, 1937.)
Tasmania ii Barred Bandicoot; Gunn's Striped Bandicoot
PERAMELES GUNNII J. E. Gray
Perameles Gunnii J. E. Gray, Ann. Nat. Hist., ser. 1, vol. 1, p. 107, 1838.
("Van Diemen's Land" [= Tasmania].)
FIGS.: Waterhouse, 1841, pi. 15; Gould, 1859, vol. 1, pi. 9; Lydekker, 1894,
pi. 21.
While this species still occurs in numbers in Tasmania, it is
"bordering on extinction in Victoria" (David H. Fleay, in litt.,
June 1,1937).
Muzzle tapering, gray-brown; under parts, feet, tail, and four
ORDER MARSUPIALIA: MARSUPIALS 49
broad bands on each side of the rump white (J. E. Gray, in Gunn,
1838, pp. 107-108). General color grizzled yellowish brown; outer
surface of ears yellowish brown, with a darker terminal blotch;
sides of rump with four more or less distinct pale vertical bands
running downward from near the middle line, the spaces between
them brown or black. Head and body, 380-400 mm. ; tail, 80-90 mm.
(Thomas, 1888, pp. 245-246.)
Its known range includes Tasmania and Victoria. In the former
state "the bandicoots are very numerous everywhere; they . . .
live principally on roots. I knew one gentleman's entire collection
of Cape bulbs, principally Babianae, eaten by them, and I suffered
considerably myself, having lost some entire species of bulbs
through these animals." (Gunn, 1838, pp. 102-103.) Gray (in Gunn,
1838, p. 108) records insect remains found in the stomach of one
specimen.
This species is to be met with throughout Tasmania, but it ap-
pears to be less commonly and evenly distributed than the Short-
nosed Bandicoot (Lord, 1928, p. 20) .
"At one time distributed through western and central Victoria,
this species is now restricted to a single locality near Hamilton and
its numbers are few" (C. W. Brazenor, in litt., March 3, 1937).
Western Barred Bandicoot; Marl
PERAMELES MYOSURA MYOSURA Wagner
Perameles myoswos Wagner, Archiv fur Naturg. (Wiegmann), 7th yr., vol. 1,
p. 293, 1841. (The type locality, not stated in the original description, is
Swan River, according to Glauert (1933, p. 23), or King George's Sound,
West Australia, according to Iredale and Troughton (1934, p. 19).)
FIGS.: Schreber, Saugthiere, suppl. 3, pi. 155 Ad, 1842; Gould, 1845, vol. 1,
pi. 10.
"No specimens have reached the [Perth] Museum since 1900. It
is therefore assumed that the animal is extinct." (Glauert, 1933,
p. 23.)
Above mixed with blackish and yellowish brown; below dirty
yellowish white; ears pale dusky, with a rusty-red spot at external
base; a dark band extending across sides in front of thighs; feet
whitish; tail scaly, short-haired, dusky above, dirty white below.
Head and body, 11 inches; tail, 3 inches. (Wagner, 1841, pp. 293-
295.)
The former range was the southwestern portion of Western
Australia. According to Gould (1863, vol. 1, p. 14), it "inhabits
the whole line of coast of the Swan River colony, but, so far as I
can learn, is not found to the westward of the Darling range of
hills." He adds that "its food consists of insects, seeds, and grain."
3
50
EXTINCT AND VANISHING MAMMALS
"Apparently not plentiful in the South-west, although described
by natives as being fairly numerous in the Salt River district. A
species of Bandicoot, probably this species, is said to have formerly
extended as far north on the mainland as Sharks Bay." Specimens
are recorded from the vicinity of Pin jelly and Kojonup. (Short-
ridge, 1910, pp. 833-834; map, p. 835.)
FIG. 6. Western Barred Bandicoot (Perameles myosura myosura}.
After Gould, 1845.
South Australian Barred Bandicoot
PERAMELES MYOSURA NOTINA Thomas
Perameles myosura notina Thomas, Ann. Mag. Nat. Hist., ser. 9, vol. 10,
p. 144, 1922. ("Plains near the head of the St. Vincent Gulf," South
Australia.)
FIGS.: Jones, 1924, p. 147, fig. 102; Le Souef and Burrell, 1926, fig. 83.
This subspecies is apparently extinct in southeastern South
Australia but is "probably holding its own in the semi-arid Nullarbor
Plain" in the west (A. S. Le Souef, in Hit., February 15, 1937) .
It closely resembles P. fasciata in coloration, with three distinct
black bands on the hind quarters; skull with more slender muzzle
and smaller teeth than in P. fasciata. Head and body, 280 mm.;
tail, 90 mm. (Thomas, 1922, p. 144.)
Formerly it seems to have ranged across the entire east-west
extent of South Australia, in the more southern parts. According to
Jones (1924, pp. 149-150), "this beautiful little Bandicoot had at
one time a fairly wide distribution in this State. In addition to the
animals from the head of St. Vincent Gulf, are others from the
River Murray in South Australia, and from Adelaide itself. As
far as can be ascertained it has now disappeared from all these
localities, and remains only in the wastes of the western portion
of the Centre. .
ORDER MARSUPIALIA: MARSUPIALS 51
"Barred Bandicoots become very tame and familiar in captivity,
but . . . they are desperately pugnacious among themselves. On
one occasion eight live specimens were sent from Ooldea. All eight
were dead . . . when they arrived in Adelaide. . . . But among the
corpses were four pouch young, which were uninjured. ... In the
end they all recovered." A female from this lot eventually bred
freely in captivity. Two young were generally found in a litter.
Rabbit-eared Bandicoot; Rabbit-bandicoot; Bilby; Dalgite;
Pinkie
MACROTIS LAGOTIS LAGOTIS (Reid)
Perameles Lagotis Reid, Proc. Zool. Soc. London 1836, p. 129, 1837. ("In
Australia Occidentali et in Terra Van Diemen." Thomas (1888, p. 225)
lists the type specimen from "Swan R., W. A.")
FIGS.: Waterhouse, 1841, pi. 12, and 1846, vol. 1, pi. 13, fig. 1; Gould, 1845,
vol. 1, pi. 7; Le Souef and Burrell, 1926, fig. 79; Pocock, 1937, p. 617, fig.
(subsp.?).
The several subspecies of this Australian animal (lagotis, cam-
brica, grandis, interjecta, nigripes, and sagitta) seem to be more or
less seriously reduced in numbers; one is apparently on the verge
of extinction, and another quite extinct. They will be treated in turn.
Concerning the group as a whole, Jones says (1924, pp. 164-167) :
By the early colonists the Bilby was not only regarded as an animal against
which the methods of the exterminator need not be employed; it was even
accorded a certain amount of protection, and was at times kept as a pet
about the house. The tolerance with which it was regarded by people whose
hands may be justly said to have been against all animals was due to the
fact that it was recognised that, in the destruction of mice and insects, it
played an extremely useful part. Unfortunately this regard for the Bilby
seems to have been forgotten by a later generation, and in more recent days
but little mercy has been shown to them by any section of the community. . . .
The reason for the rapid decrease in numbers of the Bilby is not quite
obvious. Certainly these useful animals have been ruthlessly slaughtered in
all districts within reach of the more settled areas. Their pelts have been
marketed in the skin sales in Adelaide in very large numbers; and they have
been more wantonly killed for "sport." Large numbers have been killed or
maimed in steel traps set for rabbits, and possibly many have fallen victims
to poison baits. As with all the more defenceless marsupials, the introduced
fox has probably played its sinister part. But in the Centre, where the fox
is still absent, or rare, and where the Bilby is but little molested by man, it
seems that some other factor must be invoked; and this is probably the
extraordinary abundance of rabbits, and the consequent struggle for breeding
burrows. There is certainly no part of this State [South Australia] where the
Bilby is not a rapidly disappearing animal.
Troughton remarks (1932, p. 221) : "According to Wood Jones,
one or two constitutes the usual litter [in members of this genus],
although there are eight teats, and it seems possible that a reduced
52 EXTINCT AND VANISHING MAMMALS
rate of breeding, in the less hospitable regions to which settlement
is forcing them, where the cunning introduced enemy [the fox]
probably now abounds, must ultimately lead to the extinction of this
harmless, picturesque, and pest-destroying marsupial."
In the typical subspecies, M. I. lagotis, the general color is gray;
head, neck, and back washed with chestnut; sides of body and base
of tail pale chestnut; ears long, broad, ovate; outer, upper surface's
of limbs grayish; under parts white; middle part of tail black;
terminal part white, with a crest of stiff hairs. Head and body, 18J
inches; tail, 10 inches. (Reid, 1837, pp. 129-130.) This is a large
race, with a long, silky coat; the black portion of the tail usually
longer than, or equaling, the crested white part (Troughton, 1932,
p. 227).
Its range includes south and central Western Australia and
extends to Central Australia.
Gould (1863, vol. 1, p. 11) considers it "tolerably abundant
over the whole extent of the grassy districts of the interior of the
Swan River colony." By retreating into its deep, long burrows, "it
frequently eludes the pursuit of the natives, who hunt it for the
sake of its flesh." He speaks of its flesh as "sweet and delicate,"
resembling "that of the rabbit."
Its status in Western Australia is summed up by Shortridge
(1910, pp. 832-833; map, p. 832) :
"Although widely distributed throughout the South- West (except
near the coast) , North-West, and Centre, it has within recent years
become extremely rare in the far interior. Most plentiful in the
inland districts of the South-West, rather frequently caught in traps
set for rabbits along the rabbit-proof fence. In the dry North-
Western and South-Eastern divisions, where it is rare, it extends to
the coast."
The same author states (1907, pp. 770-771) that in the interior
"it seems to have almost left parts of the country where it was onca
well known perhaps on account of the succession of droughts in-
land of late years."
Troughton (1932, p. 227) mentions specimens from Gracefield,
Coorigan, and Teuterden, Western Australia.
Glauert (1933, p. 24) records it in Western Australia as "widely
distributed . . . south of the Kimberley Division. The western limit
seems to be the Darling Range, although the Museum has odd
specimens from Perth and Upper Swan on the Coastal Plain. The
animal occurs as far south as Cranbrook and Jerramungup, near
the Stirling Range, and as far east as Gnawlbat, 126 degrees 15
minutes east, 26 degrees 21 minutes south." He writes (in litt.,
ORDER MARSUPIALIA: MARSUPIALS 53
March 17, 1937) that it is "affected by fumigation of rabbit burrows
in agricultural areas."
Finlayson (1930, p. 178; 1931, p. 161) records specimens from
north of the Musgrave Ranges, in the extreme north of South
Australia, and from two localities north of the Macdonnell Ranges
in Central Australia.
E. Le G. Troughton writes (in Hit., April 16, 1937) that it "may
survive always in parts of the Centre, but should be given total
protection in the south-west as its destruction of rats and mice
far outweighs any slight damage it may do."
New South Wales Bilby
MACROTIS LAGOTIS CAMBRICA Troughton
Macrotis lagotis cambrica Troughton, Australian Zool., vol. 7, pt. 3, p. 230,
1932. ("Bathurst," New South Wales.)
This eastern subspecies, extinct in Victoria and last recorded
from New South Wales in 1912, apparently survives in uncertain
numbers in southern Queensland.
It is about equal in size to the large western subspecies (lagotis) ;
the fur is shorter and more woolly ; upper parts more fuscous ; under
parts yellowish. Head and body, 390-500 mm.; tail, 248-278 mm.
(Troughton, 1932, p. 230.)
According to Troughton (1932, p. 230; map, p. 231), it was
"originally distributed over inland New South Wales from the
Darling River (Bourke) in the west, across to near the Great
Dividing Range in the east (Bathurst and Ghoulburn) , south to the
Murray River and north to the Queensland border (Moree) ; prob-
ably extending into southern Queensland."
Since 1892, fifteen specimens reached the Museum, making in all at least
twenty-two, of which the last was received from Moree in June, 1908; though
several are not definitely localised, there is no doubt that the entire series
came from within New South Wales. . . . Probably never very plentiful,
the rabbit-bandicoot was apparently distributed fairly evenly west of the
dividing range in the early days, and, unless mere coincidence, appears to
have been more abundant in some years as three specimens reached the
Museum in August, 1897, and again in 1903. There seems no doubt, however,
that the local race has vanished from the more settled areas, and that . . .
the Bilby is rapidly disappearing from New South Wales, or is at least faced
with a precarious existence in more desert regions. I am not aware of pelts
ever having been marketed to any extent in Sydney, . . . but no doubt num-
bers have been killed in rabbit traps, and wantonly for so-called sport, while
foxes must be contributing to the apparent annihilation within New South
Wales.
The last record of the Bilby's occurrence in New South Wales, so far as I
am aware, is [that of] a pair under close observation in the rocky hills on
the Wagga Experimental Farm for about five years prior to 1912, when
54 EXTINCT AND VANISHING MAMMALS
they were unfortunately slaughtered by shooters from the town. (Troughton,
1932, p. 220.)
In Victoria this Bilby was always confined to the northwestern
corner of the state, and the last record was in 1860 (C. W. Brazenor,
in Hit., March 3, 1937) .
Jones (1923a, p. 342) speaks of examining a living specimen from
Queensland. Finlayson states (1934, p. 229) that it is apparently
absent from the Dawson Valley, Queensland, but that it occurs at
Epping in the Clermont district, 150 miles northwestward.
Nalpa Bilby
MACROTIS LAOOTIS GRANDIS Troughton
Macrotis lagotis grandis Troughton, Australian Zool., vol. 7, pt. 3, p. 229, 1932.
("Nalpa, in the Lake Alexandrina District, south of Adelaide, South
Australia.")
Fia.: Jones, 1923a, p. 333, fig. 352.
.This subspecies is apparently extinct (Troughton, 1932, p. 230).
It is the largest subspecies; its ear, however, is proportionately
shorter than in the other subspecies. Head and body, 550 mm.;
tail, 260 mm.; ear, 77 mm.
It is known only from the "South-east of South Australia" (Ire-
dale and Troughton, 1934, p. 20) .
Before this form was distinguished from the typical lagotis,
Jones (1924, pp. 156-157) wrote of it as follows:
Thalacomys lagotis, though formerly abundant in South Australia, is now
either extinct or on the verge of extinction. It was the familiar species of
Bilby in the more fertile portions of South Australia only a comparatively
short time ago. Not more than thirty years since it was usual for rabbit
trappers, even in the immediate neighbourhood of Adelaide, to take more
Bilbies of this type than rabbits in their traps. This race . . . apparently had
its last South Australian stronghold at Nalpa and in the wide tract of
country about Lake Alexandrina; but from Nalpa it has long since disappeared,
and it seems most probable that the animal is now extinct in this State.
Rawlinna Bilby
MACROTIS LAGOTIS INTERJECTA Troughton
Macrotis lagotis interjecta Troughton, Australian Zool., vol. 7, pt. 3, p. 227,
1932. ("Rawlinna, Trans-Australian Railway, Western Australia.")
This subspecies seems to be known definitely from only two
specimens, both taken at the type locality.
^Smaller than the western lagotis; fur shorter and more woolly;
general color more drab-gray and less contrasting; under parts
whitish; black portion of tail equaling, or shorter than, the white
ORDER MARSUPIALIA: MARSUPIALS 55
terminal portion. Head and body, 303-318 mm.; tail, 207-232 mm.;
ear, 80-82 mm. (Troughton, 1932, pp. 227-228.)
"It is possible that the Musgrave Ranges specimen recorded by
Finlayson [as lagotis] is an aged female of this race" (Troughton,
1932, p. 228) .
Black-footed Bilby
MACROTIS LAGOTIS NIGRIPES (Jones)
Thalacomys nigripes Jones, Rec. S. Australian Mus., vol. 2, no. 3, p. 347,
figs. 358-360, 1923. ("Ooldea Soak," Trans-Australian Railway, South
Australia.)
FIGS.: Jones, 1923o, p. 347, fig. 358, and 1924, p. 152, figs. 106, 107.
"This animal is, so far, only known from the district round
Ooldea Soak, on the railway from Port Augusta to Perth. In that
district it appears to be by no means uncommon." (Jones, 1924,
p. 163.)
Smaller than M. I. lagotis but much like it in general color;
distinguished from all other members of the genus by its black
feet; under parts pure white; black portion of tail shorter than
terminal white portion. Head and body, 365-390 mm.; tail, 200-
220 mm.; ear, 105-110 mm. (Jones, 1923a ; pp. 347-350.)
The half-dozen known specimens were all collected by abo-
riginals (Jones, 1923a, p. 349) .
"On the Nullarbor Plain, in the state of South Australia, occa-
sional holes would be met with, the animals numbering, in 1921,
about 21 to the square mile" (Le Souef and Burrell, 1926, p. 299).
Although these authors add that the form was probably sagitta,
there would seem to be equal or greater likelihood of its being
nigripes, since the type locality of the latter borders on the Null-
arbor Plain.
Barcoo Bilby
MACROTIS LAGOTIS SAGITTA (Thomas)
Thalacomys sagitta Thomas, Ann. Mag. Nat. Hist., ser. 7, vol. 16, p. 426, 1905.
("Killalpanima [= Killalpaninna], east of Lake Eyre," South Australia.)
Although still considered "by no means uncommon" (Finlayson,
1935c, p. 233), this Bilby, like other members of the genus, is
probably seriously menaced by the predatory fox and the com-
peting rabbit.
This is smaller than any subspecies except interjecta, and a
little paler than lagotis. Head and body, 316-385 mm.; tail, 215-
245 mm.; ear, 79-84 mm. (Thomas, 19056, p. 426; Troughton,
1932, p. 229; Finlayson, 1935c, pp. 234-236.)
56 EXTINCT AND VANISHING MAMMALS
The Barcoo Bilby has been recorded in northeastern South
Australia, from Miller's Creek and Coward Springs, southwest of
Lake Eyre, to the Goyder's Lagoon area toward the northeastern
corner of the state. It also seems to range northward to the region
about Charlotte Waters, Central Australia. (Jones, 1923a, p. 344;
Finlayson, 1935c, p. 233.)
Jones writes (1924, p. 160) : "This Bilby is a northern form
living in the region of the great drainage system of Lake Eyre. It
is probable that it is still fairly abundant in those portions of this
legion where foxes have not yet become plentiful, and where it
can still compete with rabbits for nesting burrows."
On this subject Troughton says (1932, p. 221) : "My own
experience when collecting in the very dry country about Farina
[south of Lake Eyre], South Australia, in 1919, was that foxes
were very numerous and already tending towards a small lean
desert type capable of entering the larger rabbit burrows without
difficulty, and doubtless those of the Bilbies as well."
Six specimens obtained in the Goyder's Lagoon area about 1932
were, according to Finlayson (1935c, p. 233), the first ones to be
examined in the flesh since the type specimen was taken in 1903.
Certain notes on "Peragale lagotis" from the Charlotte Waters
region of Central Australia, published by Spencer (1896, p. 17, and
1897, p. 9) before sagitta was recognized, actually refer, it seems,
to the latter form (c/. Troughton, 1932, p. 233). "This is not
uncommon, judging by the number of tails used by the natives
as ornaments. They tie the white terminal tufts together in bundles
of from twelve to twenty." The animal occupies the inner end
of its burrow, and the natives secure it by digging it out.
White-tailed Bilby; White-tailed Rabbit-bandicoot
MACROTIS LEUCURA LEUCURA (Thomas)
Peragale leucura Thomas, Ann. Mag. Nat. Hist., ser. 5, vol. 19, p. 397, 1887.
("Exact locality . . . not . . . recorded.")
SYNONYM: Thalacomys minor miselius Finlayson (1932).
FIGS.: Thomas, 1888, pi. 2; Finlayson, 1935, pi. opp. p. 63.
According to Finlayson (1935c, p. 232), the probability of the
identity of M. I. leucura and T. m. miselius is very great, and the
latter name is here considered, at least provisionally, as a synonym.
Proportions and fur of leucura as in M. lagotis; general color
pale yellowish fawn; under parts pure white or yellowish white;
limbs pure white; tail slender, wholly white-haired, with a terminal
dorsal crest. Measurements of the very young type: head and
body, 142 mm.; tail, 116 mm. (Thomas, 1887, pp. 397-398.) In
miselius the central two-fifths of the tail has a median dorsal line
ORDER MARSUPIALIA: MARSUPIALS 57
of pale slate, bordered by fawn; head and body, 250 mm.; tail,
155 mm.; ear, 72 mm. (Finlayson, 1932, pp. 168-169).
The type of leucura, from an unknown Australian locality, was
described in 1887. A second specimen was taken at Mungerani,
east of Lake Eyre, in 1924. The 12 specimens on which the name
miselius was founded were taken in 1931 near Cooncherie on the
lower Diamantina River, in northeastern South Australia, at about
latitude 26 32'. In this area the animal was plentiful (Finlayson,
1935c ; p. 227). It appears to be known, however, from a total of
only 14 specimens.
A Wonkonguroo boy, who obtained most of the specimens near
Cooncherie, was adept at locating the burrows in sand hills,
although the entrances were blocked with loose sand. The animal
is evidently used as food by the natives. (Finlayson, 1935c, p. 227.)
"It now appears . . . that the ... composite species [M. I.
leucura and M. I. minor] has a wide central distribution in which
it may survive indefinitely, though the advent of the fox and
rabbit are considered by Professor Wood Jones to have already
exercised a marked influence on sub-desert populations, in asso-
ciation with prolonged dry seasons" (E. Le G. Troughton, in litt.,
April 16, 1937) .
Lesser Bilby; Lesser Rabbit-bandicoot
MACROTIS LEUCURA MINOR (Spencer)
Peragale minor Spencer, Proc. Royal Soc. Victoria, n. s., vol. 9, p. 6, pi. 2,
figs. 1-4, 1897. ("Sand-hills about forty miles to the north-east of
Charlotte Waters," Central Australia.)
This animal seems to be definitely known only from a small
series taken at the type locality in Central Australia more than
45 years ago.
Fur long and silky; general color fawn-gray; chin and inner
side of forelimbs white; rest of limbs and under parts gray; feet
white above; basal two-thirds of tail dark above; final third
white, crested; sides and ventral surface of tail white. Head and
body, 200-270 mm.; tail, 118-160 mm.; ear, 68-92 mm. (Spencer,
1897, pp. 6-7.)
"The 'Urpila' (P. minor) during the winter months lies within a
foot or so of the entrance of his [burrow]. . . . This peculiarity
is taken advantage of by the natives who jump on the surface of
the ground behind the 'Urpila' breaking it in and so cutting off
his retreat to the inner chamber. He is thus compelled to rush
out through the entrance where a native is waiting to give him
his quietus." (Byrne, in Spencer, 1897, p. 9.)
58 EXTINCT AND VANISHING MAMMALS
The fox and the rabbit have probably been decisive factors in
the depletion or disappearance of this animal.
Eastern Pig-footed Bandicoot
CHAEROPUS ECAUDATUS ECAUDATUS (Ogilby)
Perameles ecaudatus Ogilby, Proc. Zool. Soc. London 1838, p. 25, 1838. (Left
(south) bank of the Murray River, near the junction with the Murrum-
bidgee River, in Victoria. Not New South Wales, as stated by Iredale
and Troughton (1934, p. 21). Cj. Mitchell, 1838, vol. 2, p. 131.)
FIGS.: Gould, 1845, vol. 1, pi. 6 (central fig.); Jones, 1924, p. 167, fig. 124.
This unique little animal has apparently vanished from eastern
Australia and southern South Australia; possibly it maintains a
slight foothold (as one subspecies or the other) in Central Australia.
Ears long, elliptical, and nearly naked; muzzle much attenu-
ated; body about the size of a small rabbit, and the fur very
much of the same quality and color as in that animal; two toes
on forefeet, similar to those of a pig; tail [accidentally] wanting
(Ogilby, 1838, pp. 25-26). General color coarsely grizzled gray,
with a tinge of fawn; under parts white; limbs long and slender;
tail black above, gray below and on sides. Head and body, 250
mm.; tail, about 100 mm. (Thomas, 1888, pp. 251-252.)
The former range included the interior parts of Queensland, New
South Wales, and Victoria; also South Australia. The systematic
status of the Central Australian animal is apparently not settled,
but in coloration it is said by Spencer (1896, p. 17) to resemble
the western subspecies, C. e. occidentalis.
"The quaint and singularly gentle Pig-footed Bandicoot which
had been discovered by Mitchell in 1836 was reported by Krefft
twenty years later as exceedingly rare and disappearing as fast as
the native population" (Troughton, 1932, p. 188). This was due
to the increase of cattle and sheep (Lydekker, 1894, p. 148) .
The species is recorded from western Queensland by Longman
(1930, p. 64).
There were a few records from extreme northwestern Victoria
(the last one in 1857), and the animal is now extinct in that state
(C. W. Brazenor, in litt., March 3, 1937).
Jones writes (1924, p. 171) concerning the species in South
Australia :
Specimens in the South Australian Museum come from Cooper's Creek,
from near Ooldea, and from the Gawler Ranges. Probably it still lives in
the neighbourhood of Ooldea, but specimens have not been met with in
that district for some years. ... In 1920 one was killed between Miller's
Creek and Coward Springs to the south and west of Lake Eyre. . . . Although
its distribution in the Centre is wide, it has always been a very rare animal,
and now must be regarded as a disappearing one. . . .
ORDER MARSUPIALIA I MARSUPIALS 59
Pig-footed Bandicoots are said . . . when chased by dogs, to seek the
shelter of hollow logs or hollow trees. In the districts to which they are now
confined they would be hard put to find a log, let alone a tree .... Once
open country of this type has been invaded by the fox, the fate of Choeropus
is sealed. . . . The name by which it is known to the Kukata blacks is
Wilalya, and they regard it as an animal which has always been rare and
which is now extinct in their country.
Reporting on the Horn Expedition to Central Australia, Spencer
says (1896, pp. 17-18) :
"At the present time this is one of the most difficult of the smaller
marsupials to secure. . . . During the expedition we were unable
to secure a single specimen. On a subsequent visit to Charlotte
Waters I was fortunate enough to obtain one secured by the
blacks. . . .
"There is no doubt but that the range of the animal extends
widely over the central area. In the Adelaide Museum is a speci-
men from Barrow Creek, which lies well within the tropics, and
throughout the whole of our expedition all the natives were well
acquainted with it. ... It ... is evidently rapidly becoming ex-
tinct, except perhaps in the more central districts."
"Said to still have a wide but sparse distribution in the central
region, there has been little proof of late, and its terrestrial, non-
burrowing, specialized habits and frail constitution render its ulti-
mate extinction certain" (E. Le G. Troughton, in litt., April 16,
1937).
Western Pig-footed Bandicoot
CHAEROPUS ECAUDATUS OCCIDENTALS Gould
[Choeropus] occidentalis Gould, Mamm. Australia, vol. 1, p. 10, pi. 6,
1845. ("The interior" of "Western Australia"; type locality shown by
Thomas (1888, p. 252) to be "Boorda, Kirltana, W. A.")
FIGS.: Gould, 1845, vol. 1, pi. 6 (right and left figs.) ; Waterhouse, 1846, vol. 1,
pi. 13, fig. 2.
This animal is extinct, at least in Western Australia (L. Glauert,
in litt., March 17, 1937). The form that once occurred in Central
Australia (see discussion under C. e. ecaudatus) does not seem to
have been recorded for some years and may have suffered the
same fate.
The western subspecies differs from the eastern one chiefly in its
orange-brown rather than gray coloration.
Gould states (1863, vol. 1, p. 10) that Gilbert sent two specimens
from Western Australia and that the animal is confined to the
interior. According to Waterhouse (1846, vol. 1, p. 392), one of the
specimens came from the Swan River district.
"I was not able to find out anything definite about the dis-
60 EXTINCT AND VANISHING MAMMALS
tribution of this species in Western Australia. It is evidently very
rare. The specimen obtained by Gilbert in 1843 seems to have been
the only one ever secured in this State." (Shortridge, 1910, p. 835;
map, p. 836.)
"Mr. A. Le Souef states in a letter 2/12/1927 that he has seen a
dried skin at Rawlinna. This is the only recent record known to
me." (Glauert, 1933, p. 24.)
Family PHALANGERIDAE : Phalangers, etc.
This family consists of approximately 14 Recent genera and 110
forms. Its range extends from Tasmania and Australia to New
Guinea and the Admiralty and Solomon Islands on the north and
to Celebes and Timor on the west. Three Australian species are
discussed herein.
Honey Mouse; Honey Possum; Long-snouted Pouched Mouse
TARSIPES SPENSERAE J. E. Gray
Tarsipes Spenserae J. E. Gray, Ann. Mag. Nat. Hist., ser. 1, vol. 9, p. 40, 1842.
("King George's Sound," Western Australia.)
FIGS.: Gould, 1845, vol. 1, pi. 5; Waterhouse, 1846, vol. 1, pi. 11, fig. 1;
Cabrera, 1919, pi. 13, fig. 5; Troughton, 1923, pi. 23, and p. 152, fig.;
Troughton, 1924, pp. 128, 129, figs.
This rare, local, and unique little animal of Western Australia is
"becoming rarer" (L. Glauert, in Hit., March 17, 1937) .
Body mouselike; head elongate, tapering; general color blackish
gray ; back with a median black streak, bordered by a brown stripe
on each side; under parts pale bay; tail elongate, tapering, short-
haired, scaly. Head and body, 3^ inches; tail, 3 inches. (J. E. Gray,
1842, p. 40.) Tongue slender, protrusile, and brushlike, specialized
for thrusting into flowers for nectar. Head and body, 71 (male) to
86 mm. (female) ; tail, 95 (male) to 101 mm. (female). (Troughton,
1923, pp. 153-154.)
The range, according to Glauert (1933, p. 25), is "South-Western
Australia from the Irwin River south of Geraldton to the south
coast as far east as Esperance.
"Usually more or less coastal, but has been found along the
Great Southern as far north as Wagin, and at Nyabing east of
Katanning. The animal still occurs close to Perth in suitable
localities. . . . King George's Sound ... is still the headquarters
of the species."
Gould (1863, vol. 1, p. 9) recorded it "from Swan River to King
George's Sound, but from its rarity and the difficulty with which it
is procured, notwithstanding the high rewards I offered, the natives
only brought me four specimens."
ORDER MARSUPIALIA: MARSUPIALS 61
Shortridge (1910, p. 826; map, p. 827) records eight specimens
from Albany. "The small marsupial mice are very difficult to secure
on account of their rarity, and their nocturnal, arboreal, and to a
great extent insectivorous habits, being known chiefly from cats
killing and bringing them into houses."
Troughton, who has contributed most of the recent information
on the species, writes (1924, pp. 127-132) :
Alas, as settlement increases, the time seems near when there may be no
living representatives of these unique creatures to occupy the queer niche
which the process of evolution has fashioned for them within its fabric. . . .
Failure [to secure specimens on a collecting trip near Albany in 1922]
was not surprising, all accounts confirming Mr. Morgan's statement that the
mice visit areas periodically according to the flowery food supply, and that
they are but rarely seen except when brought in by cats as trophies of the
chase. . . .
A few months after my return . . . , the Honey Mice visited Mr. Morgan's
home once more and he has since sent over twenty adult mice to the Museum,
all of which were caught by his cat. . . .
As they are dependent upon the native flowers, the advance of settle-
ment with its periodical burning off, and the introduction of cats and other
enemies, in addition to native ones, must seriously threaten the future of
these marsupials ....
It is reassuring to hear from Mr. Glauert that the mice are still fairly
plentiful over an area of about 12,000 square miles, and that at the end of
1923 the West Australian Government was about to proclaim the Stirling
Ranges a sanctuary for the native fauna. Let us hope that these ranges may
prove a veritable stronghold for the Honey Mice, and that the flowers may
not miss their spring-cleaning from the brushy tongues for many generations
to come.
Troughton also says (1923, p. 155) : "Tarsipes is dependent upon
the flowers and foliage of its native districts, and as paddocks have
to be burnt off about every third year, the tiny marsupials are
literally hunted from paddock to post and prevented from settling
in any one area. Before the rapid advance of Western Australia's
settlement schemes, . . . fire and other enemies will send the Honey
Mice to join their fossil forbears in comparative oblivion, leaving
them represented only by a few museum skins and stray skeletons."
Leadbeater's Opossum
GYMNOBELIDEUS LEADBEATERI M'Coy
Gymnobelideus Leadbeaten M'Coy, Ann. Mag. Nat. Hist., ser. 3, vol. 20, p. 287,
pi. 6, 1867. ("Banks of the Bass River, in Victoria.")
FIGS.: M'Coy, 1867, pi. 6; McCoy, 1883, pi. 91; Lucas and Le Souef, 1909 ;
p. 107, fig.; Brazenor, 1932, pi. 6.
This rare species, the only known member of its genus, was based
upon two specimens collected in 1867 along the Bass River, South
Gippsland, Victoria. In 1900 another specimen, reputed to have
62 EXTINCT AND VANISHING MAMMALS
come from the same locality, was secured from a Melbourne dealer.
In 1909 the fourth known specimen was presented to the National
Museum of Victoria by A. G. Wilson; it came from Mount Wills in
East Gippsland, some 160 miles from the Bass River habitat. A fifth
specimen was presented to the same institution by F. V. Mason,
who had taken it many years previously at the edge of the Koo-
FIG. 7. Leadbeater's Opossum (Gymnobelideus leadbeateri)
Wee-Rup Swamp, about 3 miles due south from Tynong Railway
Station and only a few miles from Bass River. (Brazenor, 1932,
pp. 106, 108.)
Practically our entire knowledge of the species is based upon these
five specimens, which are in the National Museum of Victoria.
Its general appearance is much like that of the Sugar Glider
(Petaurus breviceps) , but it has no flying membrane. Color above
brownish gray to fawn-gray, with a dark brown to black median
stripe from head to sacrum; dark patches above and below ear and
about eye; ears large, nearly naked toward tip; chin and throat
dull buff; rest of ventral surface light yellowish gray; tail long,
bushy, colored like body (in one specimen the terminal half is
black) . Head and body, 169-200 mm.; tail, 168-203 mm. (Brazenor,
1932, pp. 106-108.)
ORDER MARSUPIALIA: MARSUPIALS 63
"There is much virgin scrubland in Gippsland in which the small
creature could survive. It is nocturnal in habits, and its general
resemblance to Petaurus breviceps is close enough to make its recog-
nition by evening light very difficult. When these facts are con-
sidered, the possibility of its survival is greater than might at first
be realised, and it is probable that a systematic search would re-
establish this small creature among the living animals of Victoria."
(Brazenor, 1932, p. 109.) More recently Mr. Brazenor has come
to the conclusion (in litt., March 3, 1937) that it is "probably
extinct."
"The risk of extermination for small unexploitable opossum forms
of restricted range is indicated by the fate of Leadbeater's opossum
. . . , an important phalangerid link, originally restricted to a small
area of Victoria where denudation of its limited scrub habitat
has apparently led to the animals' extinction. This unique mar-
supial is represented by barely a dozen specimens in state museums."
(Troughton, 1938, p. 408.)
Presumably Leadbeater's Opossum has been subject 'to attack by
the Domestic Cat and perhaps other introduced enemies. It is also
quite possible that many specimens have fallen victims to opossum
trappers who did not differentiate them or at any rate did not
realize their exceptional value.
Some of the numerous government reserves that have been
established in Victoria might provide sanctuary for this unique
animal if it were still extant.
Western Ringtail; Western Ring-tailed Opossum
PSEUDOCHEIRUS OCCIDENTALIS (Thomas)
Pseudochirus occidentalis Thomas, Cat. Marsupialia and Monotremata Brit.
Mus., p. 174, 1888. ("King George's Sound, W. A.")
This animal, confined to the extreme southwest of Western
Australia, is feared to be on the verge of extinction.
Color above deep smoky gray; limbs like back, but hands and
feet darker; under parts white; basal part of tail dark brown,
terminal two-fifths white, naked part below tip smooth. Head and
body, 335 mm.; tail, 310 mm. (Thomas, 1888, pp. 174-175.)
Nearly a hundred years ago Gould's collector, John Gilbert,
obtained specimens at Perth, Swan River, and King George's Sound.
More recently Shortridge (1910, pp. 827-829) collected 22 specimens
at Margaret River and Busselton, remarking that it is "chiefly
confined to the banks of rivers and swamps in the South-West;
local, and apparently disappearing in many places." Yet he con-
sidered it "fairly plentiful near the Margaret River." These few
64 EXTINCT AND VANISHING MAMMALS
records suggest that the animal was practically confined to the
South-West Division of Western Australia, from Perth southward.
(See map, Shortridge, 1910, p. 829.)
Glauert (1933, p. 24) gives its range as "lower South- Western
Australia in small isolated colonies, which suggest that the animal
is on the verge of extinction through natural causes."
No particular information is at hand concerning its enemies, aside
from Gilbert's remark (in Gould, 1863, vol. 1, p. 25) : "It ... is
often found in holes in the ground, . . . from which it is often
hunted out by the Kangaroo dogs."
Family PHASCOLARCTID AE : Koalas
The Koalas, consisting of a single genus and species, with three
subspecies, are restricted to eastern Australia. All forms come
within the scope of the present work.
New South Wales Koala; Native Bear
PHASCOLARCTOS CINEREUS CINERE(US (Goldfuss)
fypurus cinereus Goldfuss, in Schreber, Saugthiere, pis. 155 Aa, Ab, 1817;
Isis (Oken), 1819, Heft 2, p. 271. ("The forests of New Holland, about
50-60 English miles [southwest] from Port Jackson [Sydney]," New
South Wales.)
FIGS.: Waterhouse, 1841, pi. 31; Gould, 1854, vol. 1, pis. 13, 14; Lydekker,
1894, pi. 10; Nat. Geog. Mag., vol. 70, no. 6, p. 715, right-hand fig, 1936;
Pocock, 1937, p. 626, fig. (subsp.?).
Once numerous in the timbered areas of New South Wales, the
typical subspecies of this unique animal has been reduced almost to
the verge of extinction, although many thousands of the Queensland
subspecies (P. c. adustus) and perhaps a thousand of the Victorian
subspecies (P. c. victor) still exist.
The fur is dense and woolly; general color gray, either light or
dark, sometimes mottled, with whitish patches on hind quarters;
under parts, hands, and feet more or less whitish ; ears large, thickly
haired; tail rudimentary. Head and body, 700-820 mm. (Le Souef
and Burrell, 1926, pp. 291-292.) Auburn groin patches separated
by a creamy-white median area (Troughton, 1935, p. 139).
The Koala feeds almost entirely on the foliage of a few trees
of the genus Eucalyptus: E. viminalis, E. melliodora, E. rostrata,
E. microcorys, and E. maculata (Sutton, 1934, p. 78). Thus the
ranges of the three subspecies are pretty definitely restricted to
those areas in which /some or all of these eucalypts occur. The
species as a whole formerly ranged from extreme southeastern
South Australia through Victoria and the eastern half of New
South Wales into Queensland (see map, Victorian Nat., vol. 51,
ORDER MARSUPIALIA: MARSUPIALS 65
no. 3, p. 80, 1934). While the exact geographical limits of the
several subspecies have not been fully determined, we may pro-
visionally consider the range of cinereus to be New South Wales;
of adustus, Queensland; and of victor, Victoria and southeastern
South Australia.
In New South Wales Gould (1863, vol. 1, pp. 18-19) considered
the animal "nowhere very abundant" but most numerous "in the
brushes which skirt the sea side of the mountain-ranges between
the district of Illawarra and the River Clarence." He recorded it
also "among the cedar brushes of the mountain ranges of the interior,
particularly those bordering the well-known Liverpool Plains." He
prophesied that it "is certain to become gradually more scarce, and
to be ultimately extirpated."
"Though at one time extremely numerous, the koala is now, over
the greater part of its range, very scarce. This is largely due to
a disease which swept it off in millions in the years 1887-8-9, and
from 1900 to 1903. This disease took the form of ophthalmia and
periostitis of the skull. Bears are generally heavily infected with
intestinal parasites." (Le Souef and Burrell, 1926, p. 292.)
At Marrangaroo, County of Cook, N. S. W., "the native bear
was quite common then [1884-5], but quite extinct there now"
(Chisholm, 1923, p. 60) . On the Comboyne Plateau, N. S. W., it
is "very rare here now and only inhabiting the Eucalypt timber at
the edge of the Plateau" (Chisholm, 1925, p. 72).
In the fox-free eastern coastal area "there are also a few Koalas,
but these never get a chance, as the temptation to shoot or catch
the defenceless little animals as they sit exposed on a bough, is
more than the so-called sportsmen of the community can resist,
and even in our National Parks they are destroyed" (Le Souef,
1923, p. 110).
Barry writes (1928, p. 163) of the Koala's status on Kuringai
Chase, near Sydney: "Native Bears were also common here years
ago, but now, as in most places in New South Wales, they are
rarely seen."
"The typical N. S. Wales animal has been reduced to a state
verging upon extinction, in which the setting aside of adequate
areas with assured supply of favoured eucalypt diet trees presents
the only hope of survival. It is notable that any attempt to breed
them in captivity is dependent for ultimate success on the provision
of such reserves." (E. Le G. Troughton, in litt., April 16, 1937.)
Lydekker remarks (1894, p. 80) in regard to the Koala's economic
status:
The flesh is considered a great delicacy by the natives, and is regarded as
not unpalatable even by Europeans. Of its pursuit by the natives in the neigh-
bourhood of Port Jackson, Colonel Patterson writes as follows: "The natives
66 EXTINCT AND VANISHING MAMMALS
examine . . . the branches of the loftiest gum-trees, and upon discovering a
Koala, they climb the tree in which it is seen with as much ease and
expedition as a European would mount a tolerably high ladder. . . . They
follow the animal to the extremity of a bough, and either kill it with the
tomahawk or take it alive. . . ."
The Koala must be an abundant animal, since from 10,000 to 30,000 skins
are annually imported into London, while in 1889 the enormous total of
300,000 was reached. The value of these skins now ranges, according to Poland,
from five-pence to a shilling each; and they are mainly used in the manu-
facture of those articles for which a cheap and durable fur is required.
Concerning the Koala's decline, persecution, and need of protec-
tion, Jones writes (1924, pp. 184-186) :
It may be said to spend its whole life clinging to, and feeding upon, the
great eucalyptus trees. In just so much as it is a perfected specialisation
to its environment, so it is a slave to its environment. It has adapted itself to
the gum tree, and has become dependent upon the gum tree. It must be
regarded as an animal which has become phylogenetically senile as the out-
come of complete specialisation ....
Probably no animal has been so ruthlessly slaughtered in order to satisfy
the demands of the fur trade. ... In the year 1908, no less than 57,933 Koala
pelts passed through the markets of Sydney alone. That this deplorable
slaughter still goes on is evidenced by the fact that in the two years 1920
and 1921 Osborn and Anthony have ascertained that the huge total of
205,679 Koalas were killed for the fur market. Since in the fur trade Koala
pelts pass under the name of "Wombat," many people assume that the
Native Bear has ceased to be persecuted.
The complete extermination of the Native Bear would be a disgrace to
Australia, and yet, from its dependence upon a particular diet and a par-
ticular mode of life, its tenure of continued existence must always be regarded
as precarious. . . .
Horrible cruelties have been committed and recounted by those who have
slaughtered them wholesale for the sake of their pelts. Indeed, one may say,
on humanitarian grounds, that not only should the slaughter of the Koala for
the fur trade be prohibited because the animal is eminently one to protect
and not to exterminate, but it should be prohibited because, like the slaying
of seals, it is the most brutalising occupation that a human being can
undertake.
Le Souef and Burrell say (1926, pp. 291-292): "The quaint
koala . . . , perhaps, holds the affection of Australians more than
any other of their wild animals a fact for which its innocent,
babyish expression and quiet and inoffensive ways are largely re-
sponsible. It has been portrayed in caricature and verse, and its
hold on the public is used effectively by advertisers. . . .
"The skin forms a thick, serviceable fur that will stand any
amount of hard usage. Only the most callous of shooters, however,
can bring themselves to shoot such a childlike animal."
"In each of the States of Victoria, Queensland and New South
Wales, the animal is protected by law" (Stead, 1934, p. 18). Im-
portation into the United States of America was prohibited about
1930-31 by the United States Government. "So long as the United
ORDER MARSUPIALIA: MARSUPIALS
67
States market remains closed there will be but little local incentive
for destruction quite apart from any Australian protective laws"
(Wild Life Preservation Society of Australia, 22d Ann. Kept., 1931) .
Hobley calls attention (1934, p. 79) to private sanctuaries for
FIG. 8. New South Wales Koala (Phascolarctos cinereus oinereus)
the Koala established by Noel Burnet in the Pennant Hills, near
Sydney, and by C. A. M. Reid at Lone Pine, near Brisbane. (The
latter is presumably stocked with the Queensland subspecies.) He
adds:
"Great credit is due to the founders of these sanctuaries who
have been public-spirited enough to secure the safety of a number
of these creatures without any Government support. . . .
68 EXTINCT AND VANISHING MAMMALS
"The Wild Life Preservation Society of Australia is also wisely
working away with the object of establishing Koala Colonies in
such places as the Jenolan Caves Sanctuary, Lindfield Park, and
Davidson Park in New South Wales the State authorities must,
however, be persuaded to guarantee some security for the animals
established therein."
The cause of the Koala is eloquently pleaded by Troughton
(1932a, p. 192) :
"The Koala ... is utterly harmless everywhere, and what a
delight it would be for both young and old if they were plentiful
enough to haunt the suburbs and homesteads as possums often do.
They seek only the freedom of the trees, and if the continued
slaughter of such innocents leads to their extermination, it must
inevitably appear to later generations as an indictment of the
cultural degradation of our time."
Queensland Koala
PHASCOLARCTOS CINEREUS ADUSTUS Thomas
Phascolarctos cinereus adustus Thomas, Ann. Mag. Nat. Hist., ser. 9, vol. 11,
p. 246, 1923. ("0 Bil Bil, near Mimdubbera," Eidsvold, South Queensland.)
FIGS.: Faulkner, Australian Zool., vol. 3, pt. 3, pi. 16, 1923; Le Souef and
Burrell, 1926, fig. 77.
Up to 15 or 20 years ago, the Queensland Koala must have
numbered well over a million individuals; quite possibly there
were several millions. But disease and more especially the fur
trade have reduced it to a remnant of its former numbers.
It is smaller than P. c. cinereus; fur shorter; anterior back suf-
fused with dull rufous or tawny; ears far less thickly hairy, the
inner surface almost naked; under parts lighter; the prominent
groin patches rather browner and less rufous. Head and body,
600 mm. (Thomas, 1923, p. 246.)
In former times it seems to have ranged over practically all
the more southerly and easterly parts of Queensland, north to
Inkerman (lat. 19 30') and west to the Diamantina and Cooper
River basins (about long. 143). (See map, Victorian Nat., vol. 51,
no. 3, p. 80, 1934.)
"The Queensland Minister for Agriculture has said that in
1919-1930 no fewer than ... a million native bears were slaugh-
tered in Queensland. If this slaughter continues these poor animals
will be exterminated." (Gregory, 1921, p. 65.)
"Koalas . . . are now getting numerous again in Southern Queens-
land" (Le Souef, 1923, p. 109).
"In 1924, the colossal total of over two million skins of the
Koala or Native Bear were exported and mainly sold under the
ORDER MARSUPIALIA: MARSUPIALS 69
name of 'wombat' to mask the wholesale slaughter. In the Queens-
land open season of 1927, approximately 600,000 Koala were mas-
sacred by 10,000 licensed trappers." (Troughton, 1932, p. 193.)
"In Queensland Native Bears are still to be found in fair numbers,
and no doubt the Queensland Government was influenced by this
fact when it removed the protection which the animals have enjoyed
since 1919. But it is certain that even in one month their numbers
will be seriously depleted. . . . Fur and skin brokers in Brisbane
considered that before the season closed 300,000 skins would have
been disposed of. It is doubtful whether this estimated total will
have been reached, but it has to be remembered that many young
will perish when deprived of the parental care of their mothers,
which carry the little ones 'pick-a-back' from June until towards
the close of the year." (Anonymous, 1927, p. 112.)
Stead (1934, pp. 16-17) writes:
Only in a few places in Queensland are large numbers of the Koala to be
found, but only the most careful protection by the Government and by the
Australian people will prevent them from being exterminated in these places.
. . . Telling of the terrible destruction which has gone on in Queensland . . .
makes a very unhappy story, and makes one rather ashamed to think that
his own people should so cruelly destroy one of the most fascinating, harmless
and most interesting living things in the whole of the world of Nature. . . .
Very few people have any idea of the immense number of these harmless
animals killed in the one State of Queensland in only a few years before the
present season of protection was introduced. In 1927, about 600,000 were
killed during one month's open season (August), and, for the whole year,
including a so-called close season, not less than one million were slain.
Altogether, several millions of the poor little Koalas were killed in a space
of a few years in Queensland, until a great wave of public indignation put a
stop to it for the time being.
"The tenure of the koala in the Dawson Valley [Queensland]
seems to have been a 'waning one for many years, and the last open
season reduced it to such an extent that it is now a rare animal in
many parts of the valley where it was formerly very plentiful. The
process has been hastened, too, in some places, by an epidemic, and
on Coomooboolaroo in the summer of 1929 several were seen in
comatose condition at the base of feeding trees. The single example
in this condition which was examined closely was an aged male,
and though emaciated was not heavily infested with endoparasites,
nor obviously diseased organically. . . .
"It was observed .and collected at Thangool on the Cariboe, at
Coomooboolaroo, and near Mount Hedlow, on the Fitzroy." (Fin-
layson, 1934, p. 220.)
The animal now has complete legal protection in Queensland
(Stead, 1934, p. 18) .
"Perpetual universal protection is essential to its ultimate sur-
70 EXTINCT AND VANISHING MAMMALS
vival, not alone in New South Wales and Victoria where exploita-
tion, denudation of habitat, and disease have brought this unique
animal to the verge of extinction. It is in Queensland especially,
where coastal forests of the south-east provide the last stronghold
and hope of survival, that total protection should never again be
withdrawn." (Troughton, 1938, p. 408.)
Victorian Koala
PHASCOLARCTOS CINEREUS VICTOR Troughton
Phascolarctos cinereus victor Troughton, Australian Nat., vol. 9, pt. 6, p. 139,
1935. ("French Island," Western Port Bay, Victoria.)
FIGS.: Victorian Nat., vol. 51, no. 3, pis. 11-15, 1934.
Once very common over most of Victoria and in southeastern
South Australia, this subspecies had become reduced by 1934 to
about 1,000 individuals in Victoria.
Body more robust than in P. c. cinereus; coat longer, sparser,
and hairier, especially on rump and ears; coloration richer, de-
cidedly brown; ears brownish outside, white inside; auburn groin
patches extending across the inguinal region; belly brown (Trough-
ton, 1935, p. 139) . White area on throat and chest frequently pro-
longed to nape, forming a complete collar. Head and body: three
males, 800-830 mm.; one female, 730 mm. (Finlayson, 19356, pp.
223-224.)
The Koala's status in Victoria is thus reviewed by Lewis (1934,
pp. 73-74) :
There is very good evidence that forty or fifty years ago "Native Bears"
were exceedingly common over almost the whole of Victoria. Now the species
is almost extinct on the mainland, a very few Koalas surviving in the
Inverloch district and in South Gippsland around Welshpool, Toora, Foster,
etc. Others are living and, I am glad to say, thriving on the islands in
Western Port Bay. I estimate that there are now not more than 1,000 Koalas
in this State.
On the mainland of Victoria, I feel certain, the Koala is doomed to early
extinction, and will never be re-established, excepting perhaps in some reserves
which may be specially set apart for its protection and conservation, such as
the Badger Creek Sanctuary, near Healesville. . . .
From inquiries I have made among well informed people, it appears that
the favourite "sport" of the young men and boys of thirty or forty years ago
was shooting Native Bears. Their ideas of "sport" must have been very
primitive, because no more inoffensive and easily-destroyed animal than the
Koala lives in any part of the world. . . .
Immense numbers of Koalas must have been destroyed by those young
"sportsmen" of an earlier- generation than ours, but there seems never to have
been any regular hunting with a view to marketing the skins. Yet the fur
is very thick and warm, and, I am told, is in great demand by men living in
Northern Canada and Europe ....
ORDER MARSUPIALIA: MARSUPIALS 71
Apart from the shooting which so greatly reduced their numbers, I firmly
believe that the next most important factor was the bush fires which, during
the last twenty or thirty years have ravaged practically the whole of this
State. . . . The Koala falls an easy victim. . . .
Between twenty and thirty years ago, some fishermen living at Corinella
took a few Native Bears across to French Island, where . . . they thrived and
multiplied. From this island they were introduced to Phillip Island where
they are now one of the principal attractions to tourists.
Despite the drawback of practically annual fires in the scrub,
the Koalas "were holding their own on French Island until rabbits
were introduced." Cats were then liberated to cope with the latter,
but attacked the bird life, and consequently insect pests multiplied
amazingly.
"The residents, noticing the trees dying, blamed the Koalas,"
quite without justification.
"It became necessary then, in order to preserve the Koala, to
select some other place for it, and the Fisheries and Game Depart-
ment chose Quail Island, a Government reserve and sanctuary . . .
in ... Western Port Bay. To this retreat some two or three
hundred Koalas have now been transferred. ... It is hoped that
on the three islands in Western Port the Koalas will have a safe
home." (Lewis, 1934, p. 75.)
Kershaw (1934, pp. 76-77) writes as follows concerning the sanc-
tuary on Wilson's Promontory in southern Victoria:
Totally unsettled, densely timbered, and, until recent years, rarely visited
except by cattle musterers, this area has always been an ideal sanctuary.
Thirty years ago the Koala was fairly numerous in spite of the periodical
raids of skin-hunters. . . .
Following the permanent reservation of the Promontory in 1908 as a National
Park and Sanctuary for the preservation of the native fauna and flora, these
interesting animals were no longer molested .... As a result Koalas gradually
increased in numbers ....
Their immunity from interference of any kind . . . resulted in their mul-
tiplying to such an extent as seriously to threaten the existence of their
natural food plant [Manna Gum, Eucalyptus viminalis]. . . . Quite a number
of the trees had died. . . .
Action was at once taken to reduce their numbers. Where it was possible,
many were transferred to other parts of the Park, but in remote localities,
such as Oberon Bay, transport was out of the question so that it became
necessary, in order to save the remaining trees, to have a number destroyed.
[Yet] in certain localities, this particular Eucalypt was practically exterminated.
Their food-plant gone, many of the animals died, others worked back into
the more heavily timbered ranges of the interior where they found suitable
food among the Blue Gums. With a view to their acclimatization in some of
the other States several Koalas were forwarded to New South Wales, South
Australia, and Western Australia.
Native Bears are still fairly numerous in the timbered country on the
northern and eastern coasts of the Promontory and among the big timber
in the vicinity of Sealer's Cove.
72 EXTINCT AND VANISHING MAMMALS
The slight information available concerning the species in South
Australia is summarized by Jones (1924, p. 187) :
At one time the Koala was without doubt an inhabitant of South Australia,
and many men now living can remember the time when it was by no means
uncommon in certain districts of the South-eastern portion of the State. No
more than ten years ago Koalas have been killed well within the geographical
limits of South Australia. If it inhabits South Australia to-day is rather
doubtful, although reliable information would point to the fact that a
remnant of the stock may still linger not far from the Victorian border. So
far as I know no example of the South Australian race has been examined
scientifically, and no specimens seem to have been preserved. Victorian
animals were liberated on Flinders Chase, Kangaroo Island, in November,
1923, and it is hoped that they will become established in that faunal
sanctuary.
The Koala is completely protected by law in Victoria.
Family VOMB ATID AE : Wombats
The two currently recognized genera of wombats, represented by
six forms, are confined to eastern and southern Australia, Tasmania,
and islands of Bass Strait. Four subspecies are treated here.
Island Wombat; Flinders Island Wombat
VOMBATUS URSINUS URSINUS (Shaw)
Didelphis Ursina Shaw, Gen. Zool., vol. 1, pt. 2, p. 504, 1800. (Presumed by
Spencer and Kershaw (19106, p. 39) to be based upon the "Wombach" of
Hunter, in Bewick, Hist. Quadrupeds, ed. 4, p. 522, 1800. Type locality
"New Holland" = Clarke Island, Bass Strait, according to Spencer and
Kershaw (19106, pp. 37-39) ; but Cape Barren Island, Bass Strait, accord-
ing to Iredale and Troughton (1934, p. 34).)
FIGS.: Peron and Freycinet, Voyage Terres Australes, atlas, ed. 1, pi. 28,
1811, and ed. 2, pi. 58, 1824; Cabrera, 1919, pi. 17, fig. 1.
Formerly an inhabitant of several of the larger islands of Bass
Strait, this Wombat has been exterminated on all of them except
perhaps Flinders Island. It is also represented by a small colony
introduced at Eddy stone Point, Tasmania.
This is the smallest of the Wombats; hair coarse, varying from
light sandy brown to blackish; rhinarium naked. Head and body,
775 mm. Weight, 25-30 pounds (Spencer and Kershaw, 1910a, p. 29) .
This species was originally known from King, Deal, Cape Barren,
Clarke, and Flinders Islands in Bass Strait. At the time of its
discovery, about 1798, its numbers were evidently considerable.
Flinders (1814, vol. 1, p. cxxxv) found it more numerous on Cape
Barren Island than on Clarke Island; he reports it as "commonly
seen foraging amongst the sea refuse on the shore."
Home (1808, p. 304) gives an entertaining description of an
ORDER MARSUPIALIA I MARSUPIALS 73
individual secured on Flinders' voyage and kept for two years as a
pet in a house in England. It appeared intelligent as well as at-
tached to its human friends.
In their account of King Island, Peron and Freycinet (1816,
vol. 2, p. 14) describe the local Wombat as a gentle and stupid
animal, valuable for its delicate flesh. They also give an interesting
picture of its tractability. They say it had been reduced to a domes-
tic state by some English fishermen, going by day into the forests
to seek its food, and returning in the evening to the cabin which
served as its retreat.
Spencer and Kershaw (19106, p. 48) write as follows:
It is many years ago since the King Island wombat was exterminated.
When the island was visited by a party of the Victorian Field Naturalists
Club in 1887, no trace of it was discovered nor, during the process of clearing
the land that has been vigorously carried on during recent years, has any
record of a living wombat been made.
Flinders Island afforded the only prospect of securing a living specimen
of the Bass Strait species. [In 1908] a considerable part of the north, north-
east, and north-west coast line was examined, and abundant evidence was
obtained to prove that the animal, though very rare and difficult to obtain,
was not extinct. In the deserted hut of a half-caste native at Killiecrankie
two skins were found. ... On the island there are, in addition to a few
settlers, a number of half-castes .... The existence of the wombat is well-
known to them, but it is by no means easy to secure. ... On Cape Barren
Island . . . the animal was found to be quite extinct, though well-known
under the name of "badger" ....
The animal is now extinct everywhere except on Flinders Island.
An animal as large as a Wombat, always limited in numbers by
an island habitat, could scarcely be expected to survive indefinitely,
when confronted by deforestation as well as by the presence of
settlers and half-castes who evidently prized its flesh.
It is "now believed to be represented by small colonies on Flinders
Island. Observation and careful provision for their safety may be
necessary to avoid extinction." (E. Le G. Troughton, in litt., April
16, 1937.)
"The Flinders Island wombat has been introduced, and there is a
small colony ... at Eddystone Point, North-East Tasmania. They
were liberated there by the lighthouse-keepers." (Lord, 1928, p. 20.)
[The Tasmanian subspecies, Vombatus ursinus tasmaniensis
(Spencer and Kershaw), "has always and still does exist in large
numbers in Tasmania" (R. Boswell, in litt., May 13, 1937).]
[The common Wombat (Vombatus hirsutus hirsutus (Perry)) is
still more or less numerous in wild and rugged portions of south-
eastern Queensland, New South Wales, and Victoria. In southeastern
South Australia another subspecies, Vombatus hirsutus niger
(Gould), has been recognized; but no information concerning its
numerical status is at hand.]
74
EXTINCT AND VANISHING MAMMALS
Hairy-nosed Wombat
LASIORHINUS LATIFRONS LATIFRONS (Owen)
Phascolomys latifrons Owen, Proc. Zool. Soc. London 1845, p. 82, 1845. ("Con-
tinental (South) Australia.")
FIGS.: Gould, 1863, vol. 1, pis. 59, 60; Wolf, 1867, vol. 2, pi. 27; Royal Nat.
Hist., vol. 3, p. 266, fig., 1894-95; Jones, 1924, p. 267, fig. 189.
This Wombat is now practically restricted to coastal South
Australia, though once extending a little farther east and west; its
numbers have evidently been severely reduced.
FIG. 9. Hairy-nosed Wombat (LasiorhiniLs latifrons latifrons).
After Wolf, 1867.
The fur is soft and silky; general color grizzled gray, somewhat
dappled; chin dark; cheeks, throat, and chest white; belly gray;
ears long and narrow; rhinarium hairy; tail rudimentary. Head
and body, 900 mm. (Jones, 1924, pp. 266-267.)
In South Australia the species has been recorded from Mount
Gambier, Port Augusta, Port Lincoln, River Murray, River Light,
Fowler's Bay, Yorke Peninsula, Blanchetown, Blyth, 30 miles
north of Adelaide, and Nullarbor Plain. "Apparently its distri-
bution does not extend into the more northern parts of South
Australia" (Spencer, 1896, p. 3). Specimens are recorded from
Eucla, in the extreme southeast of Western Australia (Jones,
1924, p. 268). E. Le G. Troughton writes (in litt., April 16,
1937) that it was once plentiful, according to early observers,
in southwestern New South Wales and Victoria, but now is ap-
parently restricted to coastal South Australia, the inference
being that survival is not assured. A number of specimens were
ORDER MARSUPIALIA: MARSUPIALS 75
recorded by Kershaw (1909, p. 118) from Deniliquin, N. S. W.,
close to the Victorian border.
"Although in some parts of the colony [South Australia], es-
pecially on Yorke's Peninsula and about Port Lincoln, the holes of
these Wombats are very numerous, yet the animals are but rarely
seen. Many of the oldest colonists have informed me that they
never saw a Wombat alive. . . . The flesh they [the blacks] de-
scribe as being like pork, and excellent eating." (Gould, 1863, vol. 1,
p. 68.)
"It could probably hold its own under present day conditions, and
with existing introduced enemies, if only it had adequate protection
from man. That Wombats are harmless to small holders is not con-
tended. So bulky an animal which drives tunnels with such ease
is not, of course, desirable in closely settled or intensively worked
agricultural areas. But South Australia possesses vast tracts where
Wombats might burrow and live without detriment to any human
enterprise. In these areas they need protection from man alone."
(Jones, 1924, p. 270.)
"Being in grave danger of extermination, and having a distribu-
tion restricted to South Australia, it is the intention of the [Fauna
and Flora] Board to attempt to acclimatise the wombat on Kangaroo
Island; the sending of a single specimen to the Chase on October 1,
1926, may therefore be recorded; others will be forwarded as soon
as obtained" (Waite and Jones, 1927, pp. 323-324).
On the other hand, H. H. Finlayson (in Hit., March 20, 1937)
considers the species still "plentiful in a restricted habitat."
Le Souef and Burrell (1926, pp. 293, 295) write as follows:
The . . . hairy-nosed wombat is found in the drier inland areas; it also
lives along the coast of the Great Australian Bight ....
The hairy-nosed wombat has been killed out over a large part of its range.
In the Riverina, where at one time it was fairly plentiful, the settlers had
to get rid of it as part of the campaign against the rabbits, which pests had
a very secure harbour in wombat burrows.
The skin is not put to any commercial use, though the aborigines use the
fur of Ph. latijrons for making string, coils of which are wound round their
hair.
On its economic status E. Le G. Troughton remarks (in litt.,
April 16, 1937) : "Colonies were exterminated near settlement be-
cause of damage to fencing and crops, and risk of injury to stock
in the burrows."
Southern Queensland Hairy-nosed Wombat
LASIORHINUS LATIFRONS GILLESPIEI (De Vis) 1
Phascolomys gillespiei De Vis, Annals Queensland Mus., no. 5, p. 14, pis. 9, 10,
1900. (Moonie River, southeastern Queensland.)
FIG.: De Vis, 1900, pi. 10.
1 For the use of this combination, see Longman, 1939, p. 286.
76 EXTINCT AND VANISHING MAMMALS
This Wombat is "apparently now extremely rare and restricted
to remote parts of large properties." Its extinction is "apparently
imminent." (E. Le G. Troughton, in litt., April 16, 1937.)
General color gray, mixed with black, and washed with fawn
(especially on rump and back) ; inner surface of ears, throat, and
chest white; a broad curved blotch before and a spot behind the
eye, black; forearm and feet dark brown; rhinarium hairy; skull
relatively broader than in other Wombats. Head and body, 1,020
mm. (De Vis, 1900.)
The existence of a Wombat in Queensland was regarded as more
or less mythical until three specimens of the present form were
secured at the type locality and vicinity in the last decade of the
nineteenth century. The subsequent record of the animal seems
very meager.
In 1923 Wilkins (1928, pp. 25-27) made a search for it in the
Moonie River district, near Hollymount, finding "ancient tunnel-
lings of many wombats" but not encountering any of the animals.
He concluded that "there is no doubt that it is almost, if not quite,
extinct in this district."
Its range would appear to be restricted to southeastern Queens-
land.
Central Queensland Hairy-nosed Wombat
LASIORHINUS LATIFRONS BARNARDI Longman
Lasiorhinus latifrons barnardi Longman, Mem. Queensland Mus., vol. 11, pt. 3,
p. 283, 1939. ("Epping Forest Station, 75 miles west of Clermont," east-
central Queensland.)
This recently described Wombat is known from four specimens
(three of which are only skulls) , and it is considered on "the verge
of extinction" (Longman, 1939, p. 287) .
General dorsal color brown, mottled with gray, and interspersed
with black hairs; rhinarium completely clothed with short brown
hairs; ears elongate, well haired outside, with white tufts; under
parts dirty gray. Total length, 3 feet 4 inches; tail, 2^ inches.
(Longman, 1939, pp. 283, 286.)
Although Wombats "were widely distributed in Queensland in the
Pleistocene and two present-day species were known to occur spar-
ingly in southern parts of the State, it was somewhat surprising
to have definite evidence of living wombats in a locality in central
Queensland. This extends their range by over 400 miles. . . .
"Mr. Charles Barnard reports that there were many burrows in
the district, but very few tracks of the animals were seen. . . .
"Only three animals were seen, one of which was shot. As sug-
ORDER MARSUPIALIA: MARSUPIALS 77
gested by Messrs. Barnard, it is probable that these wombats were
much more numerous in earlier years, but successive periods of
drought have brought them to the verge of extinction. . . . The
specimen shot . . . has been feeding on ... stems and leaves,
including awns of the Comet River Grass, Perotis rara" (Longman,
1939, pp. 283, 286-287.)
Longman adds a report of Wombats seen distinctly about 1917
in the Tambo district, south-central Queensland.
Family MACROPODIDAE: Kangaroos, Wallabies, etc.
This largest of marsupial families contains approximately 19
genera and 125 forms. It ranges through Australia, Tasmania, New
Guinea, and neighboring islands. Accounts of 27 forms appear in
the following pages.
St. Francis Island Rat-kangaroo
BETTONGIA sp.
This extinct animal, a former inhabitant of one of the islands in
the Great Australian Bight, does not seem to be represented in the
museums by so much as a skeletal fragment upon which a technical
name might be based. Its brief and tragic history is recounted by
Jones (1924, pp. 214-215) :
Upon St. Francis Island in Nuyts' Archipelago there lived, during the
time of the present occupiers, large numbers of what was evidently a species
of Bettongia. Since the mammalian fauna of the islands of the Bight has
proved, in so many instances, to exhibit distinctions from the types inhabiting
the mainland, it is worth while recording what can still be ascertained con-
cerning this interesting and recently exterminated animal.
When the island was first settled, some forty years ago, "Rat-Kangaroos,"
or "Tungoos" were swarming. The animals do not seem to have formed
burrows, but they lived in the undergrowth, and used frequently to hop into
the homestead to take bread or other eatables thrown to them from the
table. They do not appear to have been nocturnal; they do not seem even
to have been afraid of the human invaders of the island. Their only offence
seems to have been that they had a liking for the garden produce of the
family who settled on the island.
Cats were introduced in order to exterminate the Tungoos, and their work
has been done completely. To what species the animal belonged can never
be known and the fact of its extermination in this manner is much to be
regretted.
There are many islands in the vicinity of St. Francis to which some
members of the original colony could have been transported, and so given a
chance to survive.
The story is one of importance from the point of view of legislation for
the protection of insular faunas, since it demonstrates clearly how rapidly
and how completely an interesting island fauna may be destroyed and lost
to science for ever.
78 EXTINCT AND VANISHING MAMMALS
It is much to be hoped that Isoodon nauticus, Petrogale pearsoni, Thylogale
flindersi, Leporillus jonesi, and Rattus murrayi are not permitted to follow
the Tungoos of St. Francis Island into the ranks of recently exterminated
animals.
Gaimard's Rat-kangaroo
BETTONGIA GAIMARDI (Desmarest)
Kangurus Gaimardi Desmarest, Mammalogie, vol. 2, suppl., p. 542, 1822.
(Vicinity of Port Jackson, New South Wales.)
FIG.: Quoy and Gaimard, Voy. Uranie et Phys., Zool., atlas, pi. 10, 1824
(as Hypsiprymnus white}.
This rat-kangaroo of eastern Australia is apparently extinct.
The general color is grizzled gray, with a yellowish tinge; tail
colored like body for the basal third, then darkening and the hair
lengthening until there is a distinct black crest on the terminal third ;
under side of tail white. Head and body, 390 mm.; tail, 280 mm.
(Thomas, 1888, p. 109.)
Le Souef remarks (1923, p. 110) that this is one of three mammals
that "are entirely confined to the fox area of Eastern Australia" and
"require our immediate attention if the remnants are to be saved.
... I cannot locate any Gaimard's Rat-Kangaroos; they used to
live on the Mountains and western plains of New South Wales."
Le Souef and Burrell say (1926, p. 233) : "We have noted Gaim-
ard's rat-kangaroo in the open forest on the Blue Mountains . . . ."
They add, in regard to the group of rat-kangaroos in general: "Be-
fore the advent of the fox the rat-kangaroos were extremely numer-
ous, so much so that special measures had to be taken by settlers
to protect crops and haystacks, but now many species throughout a
large part of their range are very rare, and presumably in a short
time they will be a thing of the past wherever the fox can pene-
trate."
"I think that this species is definitely extinct. I have not seen or
heard of it for upwards of 20 years." (A. S. Le Souef, in Hit.,
February 15, 1937.)
E. Le G. Troughton writes (in Hit., April 16, 1937) that it once
inhabited coastal New South Wales, but it is now apparently extinct,
possibly since the advent of the fox.
According to C. W. Brazenor (in Hit., March 3, 1937) , there are
few Victorian specimens, and the last record was in 1877.
Longman (1930, p. 59) includes southern Queensland in the range
of the species.
ORDER MARSUPIALIA I MARSUPIALS 79
Gray's Rat-kangaroo
BETTONGIA LESUEUR GRAII (Gould)
Hypsiprymnus Graii Gould, Proc. Zool. Soc. London 1840, p. 178, 1841.
("Swan River," Western Australia.)
FIGS.: Gould, 1855, vol. 2, pi. 64; Cabrera, 1919, pi. 14.
This subspecies of the Western Australian mainland has suffered
pronounced restriction of range and reduction in numbers.
Fur long and soft; general color above (including back of ears)
ashy brown; sides of head and body very faintly tinged with yel-
lowish ; under parts dirty white ; feet very pale brown ; tail brown,
except the terminal third, which is covered with longish white hairs.
Head and body, 457 mm. ; tail, 292 mm. (Gould, 1841c, pp. 178-179.)
Gray's Rat-kangaroo is apparently now confined to a compara-
tively small area in the southwest of Western Australia. Short-
ridge (1910, p. 823, map) indicates a former distribution covering
almost the entire southern half of that state. The line of demarca-
tion or intergradation between this subspecies and B. I. harveyi re-
mains undetermined.
Gould (1863, vol. 2, p. 74) "received examples of this animal from
various parts of the south-western coasts of Australia, and it appears
to be ... abundant in the plains ... in the neighbourhood of
Perth in Western Australia." He quotes Gilbert to the effect that
"it is one of the most destructive animals to the garden of the settler
that occurs in Western Australia, almost every kind of vegetable
being attacked by it, but especially peas and beans."
Thirty-five years ago it was "very abundant in many parts of the
South-West, differing curiously from the insular form in not occur-
ring near the coast." It did not then appear "to exist on the main-
land to the north of the Swan River." Specimens were recorded from
Arthur River, Woyaline Wells, Boyadine-Dale River, and Dwala-
dine. (Shortridge, 1910, pp. 822-823, fig. 258.)
More recently "this species, which was once very common in the
interior, is now confined to the Great Southern area between Beverley
in the north and Kojonup in the South" (Glauert, 1933, p. 26).
The reduction in range suggests little hope for survival unless
there exists a suitable reserve from which foxes can be excluded
(E. Le G. Troughton, in Hit., April 16, 1937) .
[On the islands of Sharks Bay, Western Australia, occurs the
typical subspecies, Lesueur's Rat-kangaroo (B. I. lesueur (Quoy and
Gaimard)), which has survived in considerable numbers, probably
owing to the protection afforded by an insular habitat. Glauert
(1933, p. 26) extends the range of this form far north along the
coast of Western Australia: "Years ago the animal was common
80 EXTINCT AND VANISHING MAMMALS
near Roebuck Bay (Broome), where K. Dahl obtained numerous
specimens."]
Harvey's Rat-kangaroo
BETTONGIA LESUEUR HARVEYI (Waterhouse)
Perameles Harveyi Waterhouse, Proc. Zool. Soc. London 1842, p. 47, 1842.
("Port Adelaide, South Australia.")
FIG.: Jones, 1924, p. 207, fig. 153.
"In certain districts it is still by no means rare, but its decrease
in numbers has been so rapid during the past twenty years that
probably the remnant still existing must not be regarded as a very
long lived one" (Jones, 1924, p. 207) .
Fur dense and soft; general color above brown, penciled with
white; sides of head and body tinged with yellowish; under parts
impure white; tail rich brown above, dirty white below, the hairs
becoming longer and white toward the tip (Waterhouse, 1842, p. 47) .
The animal is shaped like a little stoutly built kangaroo and is about
the size of a rabbit, with a short blunt head and little round ears.
Head and body, 370 mm.; tail, 300 mm. (Jones, 1924, pp. 207, 210.)
The exact distribution of this subspecies does not seem to have
been determined. It originally ranged widely through South Aus-
tralia, having been recorded from Adelaide, Port Lincoln, Gawler
Plains, Lake Phillipson, the vicinity of McDoualFs Peak, and the
northwest (Thomas, 1888, p. 113; Jones, 1924, pp. 210-211). It may
be this form that extends also into Central Australia, but the area
where it approaches or intergrades with B. L graii apparently re-
mains to be ascertained.
Gould (1863, vol. 2, p. 74) recorded it as "abundant in the plains
around Adelaide."
Spencer records "Bettongia lesueuri" from Central Australia, but
he lists no specimens and gives it the native name of "Mal-la,"
whereas Finlayson (1935, p. 62) applies the native name "maala"
to Lagorchestes hirsutus of the same general region.
Spencer's account (1896, p. 16) is as follows:
"This is the common sand-hill rat-kangaroo of Central Australia,
and is perhaps . . . the most common form of marsupial amongst
the sandy plains and sand-hills ....
"We found it during the whole course of the [Horn] expedition,
and there can be no doubt but that it is distributed right across
South, Central and West Australia."
As with so many other Australian species, we owe the chief
account of this animal's status and life history to Jones (1924, pp.
210-211) :
This Rat Kangaroo, which is probably the only living representative of
the Sub-family left in South Australia, is still existing in some numbers in
ORDER MARSUPIALIA: MARSUPIALS 81
certain districts in the North- West. Here it lives in company with the
rabbits, sharing the larger warrens with them .... The choice of a warren
seems largely to be determined by the quantity and nature of the herbage
in the neighbourhood, for in these waterless districts Rat Kangaroos are
dependent on the succulent sand hill vegetation. Rabbits are so universally
spread over the country that there probably does not exist to-day a Bettongia
colony living in its own burrows. It has thrown in its lot with the rabbit,
and although it appears to have its own appartments [sic] in the complicated
system of the large warrens, it is merely a tenant, forming a part of a
community in a manner which is rather remarkable when its exceedingly
pugnacious character is considered. Nevertheless, though it lives in apparent
harmony with the rabbits, and avails itself of the shelter of their burrows,
it is suffering for the partnership. The remnant of the Tungoos is living in an
environment in which there is a severe competition for succulent food. In
good seasons there is enough juicy herbage for cattle and rabbits as well as
Rat Kangaroos but in bad seasons the rabbits and the marsupials perish
in large numbers. Such losses among the rabbits are soon made good, but
with the marsupials this is not the case, and probably the end of the Tungoo
is not far off. When times are bad, and when the cattle and rabbits have
eaten all the herbage of the sand hills, the Tungoos become extremely bold,
and will enter a homestead in their search for anything to eat. They will
come into a room and boldly face a cat in order to obtain some potato peelings ;
they will scramble over a paling fence four or five feet high in order to get
at the vegetable garden. They are bold and enterprising little animals which
have made, and are making, a brave struggle against what seems an almost
inevitable extermination. In the more cultivated districts of the South,
where food is in plenty, the wholesale scattering of poisoned pollard has led
to their complete extinction. The poison cart has done its deadly work on
the slowly-breeding Tungoo, although the rapidly-breeding Rabbit has sur-
vived the ordeal. In the North they are steadily being pressed out of exis-
tence by the competition for food.
When we remember that their numbers in rabbit warrens, even near to
towns, was a source of constant annoyance to rabbiters less than twenty
years ago, we can realise how destructive to the native herbivorous fauna the
wholesale spreading of poisoned grain has proved to be. Nor must we forget
that the remnant which still struggles on in the North is now exposed to the
ravages of the fox.
Concerning the introduction of this rat-kangaroo on Kangaroo
Island, South Australia, Waite and Jones say (1927, p. 323) :
"Specimens bred and reared in captivity in Adelaide were liberated
within the observation enclosure on the reserve and seem to be
doing well. If, when they are turned out into the larger world, they
can avoid the goana (Varanus) they should prosper."
H. H. Finlayson (in litt., March 20, 1937) regards the species as
a whole as common in Western Australia, the Center, and north-
western South Australia.
82 EXTINCT AND VANISHING MAMMALS
Brush-tailed Rat-kangaroo
BETTONGIA PENICILLATA PENICILLATA J. E. Gray
Bettongia penicillata J. E. Gray, Mag. Nat. Hist. (Charlesworth), vol. 1, p. 584,
1837. (No type locality was stated in the original description, but Thomas
(1888, p. Ill) lists the type specimen from "New South Wales.")
FIGS.: Gould, 1841, pi. 14; Waterhouse, 1846, vol. 1, pi. 7, fig. 1; Gould,
1852, vol. 2, pi. 61; Lydekker, 1894, pi. 9 (ssp.?).
This eastern Australian subspecies is either very rare or extinct.
(The two other recognized subspecies likewise come within the scope
of the present report.)
The general color is ashy brown, penciled with white and brownish
black ; cheeks and throat faintly tinged with yellowish ; under parts
dirty white; hands and feet pale brown; tail brown above, pale
brown below, the apical third with a black dorsal crest. Head and
body, 343 mm.; tail, 285 mm. (Waterhouse, 1846, vol. 1, p. 213.)
The former range extended from the Dawson Valley, Queensland,
to Victoria, but apparently only on the inner side of the coastal
ranges.
Most of the information on this animal comes from Gould (1863,
vol. 2, p. 71), who had opportunities of studying it while it was
still abundant. "The eastern parts of Australia, particularly the
districts on the interior side of the ranges of New South Wales,
constitute the true habitat of the species .... I observed it to be
very abundant on the Liverpool Plains, and on the banks of the
river Namoi, from its source to its junction with the Gwydyr; but
between the ranges and the coast I did not meet with it." He adds
that the natives rarely pass without detecting its grassy nest on the
ground, and almost invariably kill the sleeping inmates by dashing
their tomahawks or heavy clubs at it.
It is "apparently not now found in Eastern Australia" (A. S. Le
Souef, in Hit., February 15, 1937) . "It is now very rare or extinct
in New South Wales and Victoria" (E. Le G. Troughton, in Hit.,
April 16, 1937).
C. W. Brazenor (in litt., March 3, 1937) considers the animal
extinct in Victoria, where the last record dates from 1857. There
are few Victorian examples in the National Museum of Melbourne.
According to Finlayson (1931, p. 89), "Bettongia penicillata was
taken by Lumholtz on Coomooboolaroo [in the Dawson Valley,
Queensland], but has now apparently quite disappeared from there,
and is not known elsewhere in the valley."
While no specific information concerning the causes of the dis-
appearance of this rat-kangaroo seems to have been offered, prob-
ably the fox is largely responsible.
ORDER MARSUPIALIA: MARSUPIALS 83
Gould's Rat-kangaroo
BETTONGIA PENICILLATA GOULDII Waterhouse
Bettongia Gouldii J. E. Gray, List Specimens Mammalia Brit. Mus., p. 94,
1843 (nornen nudum). ("Head of Gulph St. Vincent," South Australia.)
Bettongia Gouldii Waterhouse, Nat. Hist. Mammalia, vol. 1, p. 219, 1846.
("South Australia.")
"As far as can be ascertained at present, this animal seems to
have disappeared from South Australia" (Jones, 1924, p. 212). As
far as known, it was confined to this state.
The type specimen, a very immature animal, is the only one of
this subspecies that seems to have been described in detail. The fur
is brownish, penciled with black and yellowish white; under parts
white, more or less suffused with yellow; tail rusty brown at base,
the terminal half black both above and below (Waterhouse, 1846,
vol. 1, p. 219). Head and body, 390 mm.; tail, 310 mm. (Thomas,
1888, p. 111).
Jones (1924, pp. 212-214) furnishes practically all the available
information on the former and present status of this rat-kangaroo:
It is possible that it may prove to be still living somewhere in this State,
and if there is any hope of such a survival it would seem that the South-East
or the extreme North-East holds out the greatest promise.
Not only does it appear to have died out completely over the greater portion
of the State, but no specimen of the South Australian form seems to have
been preserved in the zoological collections in Australia. At present, so far
as this State is concerned, the race is represented only by some half-dozen
skulls. . . .
Only a few years ago this animal was extremely common over the greater
part of South Australia. Twenty years ago the dealers in Adelaide did a
great trade in selling them by the dozen at about ninepence a head for coursing
on Sunday afternoons. It may surprise people who remember those days to
know that there is not a preserved specimen, not even a skin of the animal,
available for scientific study in South Australia to-day. In the same way
it will one day surprise the rising generation when they realise that the few
native animals they are now familiar with are gone for ever. . . .
It is much to be hoped that, should some remnant of the South Australian
race be found still living in the more bush-covered portions of the South
or of the North-East, steps will be at once taken that it may be preserved
and protected by every possible means.
Ogilby's Rat-kangaroo
BETTONGIA PENICILLATA OGILBYI (Waterhouse)
Hypsiprymnus Ogilbyi Waterhouse, Naturalists' Library (Jardine), vol. 11,
Marsupialia, p. 185, 1841. ("Western Australia in the neighbourhood of
Swan River"; Thomas (1888, p. Ill) lists the type specimen from "York,
W. A.")
FIG.: Gould, 1852, vol. 2, pi. 62.
Although very plentiful in the southwest of Western Australia a
generation ago, this subspecies is now "reduced in numbers" (L.
84 EXTINCT AND VANISHING MAMMALS
Glauert, in Hit., March 17, 1937) , and concern may well be felt as
to the possibility of its following the other two subspecies on the
road to extinction.
It differs from B. p. penicillata in its generally darker coloration;
in the rusty red of the base and sides of the tail; in the rufous
coloring of the feet; in the terminal half of the tail being black
both above and below ; and in the longer ears and more slender tarsi
(Water-house, '1841, p. 186; Gould, 1863, vol. 2, p. 72). Head and
body, 360 mm.; tail, 310 mm. (Thomas, 1888, p. 111).
Gould (1863, vol. 2, p. 72) quotes Gilbert's notes from Western
Australia: "This species appears to be equally abundant in all
parts of the colony, but to evince a preference, perhaps, for the
white-gum forests. . . . This animal is one of the favourite articles
of food of the natives, who are very quick in detecting the nest, and
generally capture the little inmate by throwing a spear through the
nest and transfixing it to the ground, or by placing the foot upon
and crushing it to death."
Shortridge (1910, pp. 821-822, map) found it "very plentiful in
the South -West, where, unlike Bettongia lesueuri, it occurs near the
coast, extending as far north as the Moore River, becoming very
rare at its northern limit. Formerly recorded from Sharks Bay,
as so many of the other South-Western marsupials have been.
"Although getting scarce in the more settled districts, both species
of Bettongia are sufficiently numerous in many places to be rather
destructive to crops, on which account they are often trapped and
poisoned off in large numbers."
Shortridge records specimens from King River, Dwaladine, Woy-
aline Wells, Yallingup, and Burnside. From Perth southward, ac-
cording to Glauert (1933, p. 26), it "is found in the coastal area as
well as inland to the Great Southern and beyond." E. Le G. Trough-
ton remarks (in litt., April 16, 1937) that "survival there may be
significant of the influence of the fox not yet being fully asserted."
Rufous Rat-kangaroo
AEPYPRYMNUS RUFESCENS (J. E. Gray)
Bettongia rujescens J. E. Gray, Mag. Nat. Hist. (Charlesworth), vol. 1, p. 584,
1837. (Type locality not stated in original description, but Thomas (1888,
p. 104) lists the type specimen from "New South Wales.")
FIGS.: Gould, 1841, pi. 13; Gould, 1855, vol. 2, pi. 65; Le Souef and Burrell,
1926, fig. 51.
Once common over much of eastern Australia, this species has
largely or entirely disappeared from Victoria and New South Wales,
but it remains common in the Dawson and Fitzroy Valleys, Queens-
land.
ORDER MARSUPIALIA: MARSUPIALS
85
The fur is long and coarse; color above grizzled rufescent gray;
an indistinct white stripe crossing the sides just in front of hips;
under parts grayish white; ears rather long, black on outer surface;
tail thickly haired, pale gray above, white below. Head and body,
520 mm.; tail, 380 mm. (Thomas, 1888, pp. 103-104.)
Gould writes (1863, vol. 2, p. 75) : "The south-eastern portion of
the continent is its true habitat; and it is almost universally dis-
FIG. 10. Rufous Rat-kangaroo (Aepyprymnus rufescens)
persed over New South Wales, both on the sea and interior side of
the mountain ranges. I found it very abundant on the stony sterile
ridges bordering the grassy flats of the Upper Hunter, and in all
similar situations. . . . From its invariably seeking shelter in the
hollow logs" when startled from its nest, it "easily falls a prey to the
natives, who hunt it for food."
Of its status on the Comboyne Plateau, New South Wales, Chis-
holm says (1925, p. 73) : "Not here now, but I am informed by an
early settler that years ago they were a great pest to the farmers,
and had to be persistently poisoned. This animal appears to be fast
becoming extinct, probably largely due to the depredations of the
86 EXTINCT AND VANISHING MAMMALS
fox, as living their daylight hours in a nest on the ground they
become an easy prey for this animal."
In Victoria, according to C. W. Brazenor (in Hit., March 3, 1937) ,
the Rufous Rat-kangaroo was "once fairly common. Now rare but
probably survives in more inaccessible parts of eastern Victoria.
Last record 1905." It is completely protected by law in that state.
Finlayson writes (1931, pp. 85-86) of its status in Queensland:
This interesting animal, though highly characteristic of coastal Queensland,
has received very little mention in recent years, and there has been no pub-
lished data from which one might estimate its position in the fauna of that
State.
Strangely enough it was not taken by Lumholtz, though it must have oc-
curred in many of the districts in which he worked .... It has been twice
recorded from North Queensland by O. Thomas . . . , and by Lonnberg and
Mjoberg from Carrington . . . , but without comment, and as each record was
based on a single individual, it might be inferred to be comparatively rare.
In the Dawson and Fitzroy Valleys, however, this is far from being the
case, and it is widely spread over the whole area from sea level to the tops
of the plateaus. It occurs in almost all types of country, both open and
forested .... The banks of creeks and river flats are favourite resorts, and
there are few such places which by systematic beating cannot be made to
yield up a few. . . .
Like most of the coastal species it has little resistance to drought, and will
go to great lengths in excavating holes in dry creek beds to get down to water
level. In January, 1929, the Cariboe Creek ceased to run at Thangool, and
for miles the sandy bed thus exposed was criss-crossed with the pads of
Aepyprymnus coming down at night to drink at pot holes of their own making.
In the cattle country it is stated by squatters to have diminished considerably
in recent years, and by them it is regarded with indifference. But round
many of the newly-formed cotton settlements in The Callide Valley it is
plentiful, and at Thangool and Biloela and other points on The Cariboe has
become an unmitigated nuisance and is cordially detested by the struggling
settlers. Its raids on the crops are determined and resourceful, and as no
ordinary fence will bar them for long, poisoning is the only effective check.
Scores of thousands have been killed in this way, and skeletons (few and
far between in Museums) are littered thickly round the cotton plots.
On the outlook for the preservation of the Rufous Rat-kangaroo,
E. Le G. Troughton writes (in litt., April 16, 1937) : "There has
been a marked shrinkage of the once abundant species in coastal
N. S. Wales and Queensland, suggesting that this small, specialized,
and rather open country species is unlikely to survive, except pos-
sibly in northern coastal Queensland where it may favour less open
country, and the fox may not become established."
Gilbert's Rat-kangaroo
POTOROUS GILBERTII (Gould)
Hypsiprymnus Gilbertii Gould, Mon. Macropodidae, pt. 1, text to pi. 15, 1841.
("King George's Sound," Western Australia.)
FIGS.: Gould, 1841, pi. 15; Gould, 1854, vol. 2, pi. 69.
ORDER MARSUPIALIA: MARSUPIALS
87
The annals of this species are brief and tragic. It was discovered
in Western Australia in 1840 by John Gilbert and is represented by
his two specimens in the British Museum, but it has never since
been encountered in the flesh by a zoologist and is undoubtedly
extinct.
General color above mingled gray, brown, and black; central and
lower part of back washed with reddish brown; a blackish median
line from nose to forehead; under parts grayish white; tail black,
thinly clothed with short hairs. Total length, 558 mm. ; tail, 158 mm.
(Gould, 1841, pt. 1, text to pi. 15.)
FIG. 11. Gilbert's Rat-kangaroo (Potorous gilbertii)
Gould (1863, vol. 2, p. 79) quotes Gilbert's field notes as follows:
This little animal may be said to be the constant companion of Halmaturus
brachyurus, as they are always found together amidst the dense thickets and
rank vegetation bordering swamps and running streams. The natives capture
it by breaking down a long, narrow passage in the thicket, in which a number
of them remain stationed, while others, particularly old men and women,
walk through the thicket, and by beating the bushes and making a yelling
noise, drive the affrighted animals before them into the cleared space, where
they are immediately speared by those on the watch: in this way a tribe of
natives will often kill an immense number of both species in a few hours. I
have not heard of the Hypsiprymnus Gilberti being found in any other part
of the colony than King George's Sound.
Shortridge (1910, pp. 824-826, map) gives the following account:
"It is quite possible that they [P. gilbertii and P. platyops] are
now entirely extinct, although I picked up six old skulls of Potorous
88 EXTINCT AND VANISHING MAMMALS
gilberti near the entrances of some caves in the Margaret River
district, and they may still exist sparingly in that and other locali-
ties, as they are very liable to be overlooked on account of their
great external resemblance to Macropus brachyurus.
"The animal known to natives as 'Wurrark' around the Margaret
River is probably Potorous gilberti, said to frequent marshy country,
and although formerly numerous, it is thought to have almost, if
not entirely, died out. A few may still occur towards Cape Leeuwin."
L. Glauert (in litt., March 17, 1937) considers the species extinct.
Broad-faced Rat-kangaroo
POTOROUS PLATYOPS (Gould)
Hypsiprrymnus platyops Gould, Proc. Zool. Soc. London 1844, p. 103, 1844.
("Swan River," Western Australia; the type, according to Waterhouse
(1846, vol. 1, p. 232), is labeled as coming from "Walyema Swamps, about
forty miles north-east of Northam, Western Australia.")
FIGS.: Gould, 1851, vol. 2, pi. 70; Le Souef and Burrell, 1926, fig. 55.
This Western Australian species, regarded as rare at the time of
its discovery a century ago, and represented by apparently no
more than three specimens all told, is evidently extinct.
Face extremely broad and, with sides of body, brownish gray;
back reddish brown; face and upper surface beset with numerous
long yellowish-white hairs; under parts and limbs buffy gray; tail
brown above, paler beneath. Total length, 482 mm.; tail, 177 mm.
(Gould, 18446, p. 103.)
"This species ... is so rare that an adult male in my own col-
lection and another in that of the British Museum, both procured
fin 1840] by Mr. Gilbert in Western Australia, one in the Walyema
Swamps, near Northam in the interior, and the other at King George's
Sound, are all the examples that have yet been seen" (Gould, 1863,
vol. 2, p. 80) .
"A single specimen from the Margaret River was sent to the
London Zoological Society in 1908. This suggests that the species*
still exists in that area" (Glauert, 1933, p. 26).
Shortridge wrote in 1910 (p. 826) : "A small gregarious wallaby
is said to have been at one time plentiful in the coastal scrub to
the east of Albany; from the description it was probably one of
these species [P. platyops and P. gikbertii]. It was known to the
natives as 'Moort/ and according to them has entirely disappeared
there. Described as being rather similar to Macropus brachyurus in
habits, but more sluggish in its movements, on which account cats
and bush-fires have probably caused its disappearance."
A possible clue to the identity of the above-mentioned species
appears in Gould's original description of platyops (18446, p. 103),
wherein he cites the native name of "Mor-da," presumably current
ORDER MARSUPIALIA: MARSUPIALS
89
in the Walyema Swamps area. This bears a plausible similarity
to the "Moort" of Shortridge, whereas the native name of gilbertii
was "Grul-gyte" (Gould, 1841, text to pi. 15) or "Ngil-gyte" (Gould,
1863, vol. 2, p. 79).
For some years past the Broad-faced Rat-kangaroo has been
considered possibly or probably extinct (Shortridge, 1910, p. 825;
Le Souef and Burrell, 1926, p. 237; A. S. Le Souef, in litt., February
15, 1937; E. Le G. Troughton, in litt., April 16, 1937). Finally, L.
Glauert (in litt., March 17, 1937) definitely lists it as extinct.
FIG. 12. Broad-faced Rat-kangaroo (Potorous platyops). After Gould, 1851.
"Common" Rat-kangaroo; Long-nosed Rat-kangaroo; Dark
Rat-kangaroo; Potoroo
POTOROUS TRIDACTYLUS TRIDACTYLUS (Kerr)
Didelphis tridactyla Kerr, Anim. Kingdom of Linnaeus, p. 198, 1792. (Based
upon the "Kanguroo Rat" of Phillip, Voy. Botany Bay, p. 277, pi. 47,
1789; type locality, "New South Wales.")
FIGS.: Waterhouse, 1841, pi. 16, and Gould, 1854, vol. 2, pi. 67 (as Hypsiprym-
nus murinus); Lydekker, 1894, pi. 8; Le Souef and Burrell, 1926, figs. 53,
54; Finlayson, 1935a, pi. facing p. 99.
Formerly ranging from South Australia through Victoria and
New South Wales to southern Queensland, the Potoroo has become
extinct in South Australia and possibly in New South Wales. Its
status in Queensland does not seem to be very definitely known,
but it survives in some numbers in certain districts of Victoria.
The Potoroo is distinguished from other species of its genus by its
elongated head and short tarsus; the fur is long, loose, and slightly
90 EXTINCT AND VANISHING MAMMALS
glossy; general color dusky brown, penciled above with black and
pale brownish yellow; naked part of rhinarium extending farther
back than in P. platyops; under parts dirty yellowish white; tail
clothed with short, stiff, black hairs, extreme tip white. Head and
body, 393 mm.; tail, 235 mm. (Waterhouse, 1846, vol. 1, pp. 224-
225.)
The brief history of the species in South Australia is discussed
by Jones (1924, pp. 217-218) :
The name "common" Rat-Kangaroo although that used in all books dealing
with the marsupials, is a sadly inappropriate one. . . .
Of the former distribution of this animal in South Australia no details can
now be obtained. Save the bare record of its existence in this State [on the
Murray River], which is given in the British Museum catalogue of 1888
and which has been copied into all subsequent works, I know no other
reference to the creature as a South Australian animal. . . . The remaining
Potoroos should be carefully protected in those places where they still sur-
vive, and efforts should be made for turning them down in properly safe-
guarded sanctuaries. If this is not done there seems to be no doubt that
the remnant of the stock will share the fate of the South Australian form and
rapidly become extinct.
Finlayson writes (19356, p. 221) concerning its status in Victoria:
Few animals have been so obscure as to their status on the mainland as
the Potoroo. Its former presence in the south-eastern district of this State
[South Australia] is attested ... by the accounts of settlers, and by occa-
sional bone fragments in cave deposits, but it does not seem to have been a
common form west of the Glenelg [a river of southwestern Victoria], at the
time of settlement.
In Victoria, though better known than here, there have been few explicit
references to it in the literature, which would enable one to judge as to how it
was faring in the struggle for survival, until Mr. Brazenor, in 1933 stated that
"though very uncommon it still persists ... in the north-eastern district, in the
Grampians, and probably in the Otway Ranges," and he has since confirmed
its presence in the last locality by personally collecting it there.
I am able to add two other localities to these, viz., French Island in Western
Port, and the Portland area in the western district, and to state that in the
latter, at least, it is still plentiful. Its apparent scarcity is due, I believe,
largely to its choice of dense undergrowth .... In 1927 a rabbit trapper,
near Gorae, stated that he took over twenty of these "bandicoots" in a
short season, and this I was able subsequently to prove, by overhauling the
skulls at his dumps .... In the summer of the following year I took it
myself near Heywood and had further reports of it, and again in the winter
of the same year on French Island, and that no disaster has overtaken it
since then is vouched for by several correspondents, and very recently (for
the western district) by Professor Wood-Jones (in litt.}.
C. W. Brazenor writes (in litt., March 3, 1937) that it was once
common in eastern and southern Victoria but is now confined to
small numbers in southwestern Victoria. He adds that it is com-
pletely protected by law.
Gould (1863, vol. 2, p. 77) gives an account of it under the name
Hypsiprymnus murinus: "It is only in the swampy and damp parts
ORDER MARSUPIALIA: MARSUPIALS 91
of the brushes of New South Wales that the H, Murinus is to be
found in any abundance. The district of Illawarra, Botany Bay,
the low scrubs bordering the rivers Hunter, Manning, and Clarence,
are the principal localities in which it may be successfully sought
for."
E. Le G. Troughton (in litt., April 16, 1937) refers to the main-
land race as once common in the damp coastal regions of New
South Wales, Victoria, and South Australia, but as now rare, no
specimens having been received at the Australian Museum since
1913.
Le Souef and Burrell remark (1926, p. 233) that the Potoroo, like
all the members of the subfamily Potoroinae, makes for a hollow
log when disturbed, and thus is often easily captured.
Longman (1930, p. 59) records the species from southern Queens-
land. Finlayson (1931, p. 89) did not find it in the Dawson Valley
in that state.
[The Tasmanian subspecies, P. t. apicalis (Gould) , remains com-
mon in many localities (Lord, 1928, p. 19). Absence of the fox in
Tasmania may render that country the only hope for the survival
of any representative of the genus (E. Le G. Troughton, in litt.,
April 16, 1937).]
Desert Rat-kangaroo; Plain Rat-kangaroo
CALOPRYMNUS CAMPESTEIS (Gould)
Bettongia campestris Gould, Proc. Zool. Soc. London 1843, p. 81, 1843. ("South
Australia.")
FIGS.: Gould, 1851, vol. 2, pi. 66; Finlayson, 1932, pis. 7, 8, and 1935, pis.
facing pp. 97. 98.
"His [Finlayson's] rediscovery of the living Caloprymnus was a
romance of modern zoology. The great John Gould had received
three specimens from somewhere in South Australia in 1843. These
three specimens in the British Museum remained unique. Calo-
prymnus seemed to be as dead as the Dodo: and then Finlayson,
with the assistance of Mr Reese of Appamunna, produced [in 1931],
as a conjurer from his hat, living specimens of the long lost Plafti
Rat-kangaroo." (Jones, in Finlayson, 1935a, p. 8.)
Under fur dense and soft, very pale yellowish brown, the hairs
tipped with sooty brown; interspersed with the under fur (especially
on the back) are many long brownish white hairs, tipped with
blackish; sides dirty yellowish; under parts dirty white; feet and
tail very pale yellowish brown. Head and body, 400 mm.; tail,
355 mm. (Gould, 1843, p. 81.)
"Imagine a little animal about the bulk of a rabbit, but built
like a kangaroo, with long spindly hind legs, tiny forelegs folded
92 EXTINCT AND VANISHING MAMMALS
tight on its chest, and a tail half as long again as the body but not
much thicker than a lead pencil, and you have it in the rough. But
its head, short and blunt and wide, is very different from that of
any kangaroo or wallaby, and its coat is uniformly coloured a clear
pale yellowish ochre exactly like the great clay-pans and flood
plains." (Finlayson, 1935a, p. 102.) Head and body, 254-282 mm. ;
tail, 307-377 mm. (Finlayson, 1932, p. 165).
The exact locality from which Gould's original specimens came
(through Sir George Grey) is not now ascertainable; he merely
stated (1863, vol. 2, p. 76) that "the stony and sandy plains of the
interior of South Australia partially clothed with scrub are its
native habitat." Only the recent range of the animal can be given
with any precision. "Its proved distribution may be extended over
a large area of the eastern portion of the Lake Eyre Basin, speci-
mens and reliable records from observers personally known to the
author having been obtained from as far south as Lake Harry and
as far north as Coorabulka in South-West Queensland. The north
and south limits of its range, as at present ascertained, are, there-
fore, approximately, lats. 23 40' and 29 21' south. No records
have so far been obtained west of Lake Eyre and the Kallakoopah,
and the furthest easterly occurrence is at Innamincka, on the Barcoo,
in long. 140 49' east." (Finlayson, 1932, p. 148; map, p. 149.)
The recent history of the species may be summarized in Finlay-
son's own words (1932, pp. 150-165) :
In ordinary years the Lake Eyre Basin is a most unattractive area from
the point of view of the mammal collector, and the disappearance of
Caloprymnus from scientific ken must be attributed rather to lack of systematic
collecting than to any sudden change in the status of the animal in the
fauna, following Grey's discovery. All the evidence obtained by questioning
blacks goes to show that in all probability it has had an uninterrupted tenure
of the country, but it is equally certain that in normal times its numbers
are small, since men like Mr. Reese, whose opportunities for observation are
practically continuous throughout the year, affirm that in thirty-five years
they have seen no more than twenty specimens. . . .
At the time of my passage through the area, conditions as they bear upon
animal life were very favourable and quite supernormal as compared with
average conditions over a series of years. A period of seven years of drought
had been broken, and vegetation had been restored on a comparatively
lavish scale. All species of mammals were undergoing a quick increase in
numbers, and rodents especially had assumed plague proportions. Most of the
specimens of Caloprymnus were obtained, and the bulk of the observations
upon it were made, on two flats lying east and west of Cooncherie Sandhill. . . .
The numbers occurring in this particular locality were very considerable.
In the course of a week's riding on the two flats over an area of perhaps 20
square miles, 17 Oolacuntas were sighted. ... All the evidence obtained
so far goes to show that its distribution at present is highly discontinuous
but that it follows in a general way the fringes of the gibber plains ....
The accounts of its feeding habits given by the blacks, and several other
ORDER MARSUPIALIA: MARSUPIALS 93
items of evidence, however, would point to its being largely phytophagous,
or at least less rhizophagous than Bettongia, Potorous, and Aepyprymnus. . . .
Where Diprotodon failed [to survive], Caloprymnus may yet succeed, but
all the evidence of its physical structure is not more eloquent of changed
conditions [from relatively humid to an arid climate] than its pathetic
clinging to its flimsy grass nest, in a fiery land where a fossorial habit has
become the main factor in survival.
The first specimen flushed by Finlay son's party was run down
with a relay of horses after a chase of 12 miles. Others were taken
in the same way, while a native captured two by hand after stealing
up to their grass nests.
E. Le G. Troughton writes (in litt., April 16, 1937) : "The fact
that the otherwise defenceless animal is peculiarly suited to ex-
tremely barren and remote desert areas may ensure survival and
prevent exploitation as a rarity, but the spread of the fox, seen
personally near Marree in 1920, use as food by the blacks, and
variable seasons may continue range shrinkage to extinction."
"Common" Hare-wallaby; Brown Hare-wallaby
LAGORCHESTES LEPORIDES (Gould)
Macropus Leporides Gould, Proc. Zool. Soc. London 1840, p. 93, 1841. ("In-
terior of Australia"; according to Thomas (1888, p. 84), the cotypes
are from the "interior of New South Wales.")
FIGS.: Gould, 1841d, pi. 12; Qould, 1859, vol. 2, pi. 57; Royal Nat. Hist.,
vol. 3, p. 246, fig, 1894-95.
This species is "apparently doomed to extinction" in its last
stronghold in New South Wales (E. Le G. Troughton, in litt., April
16, 1937).
It resembles the Common Hare of Europe in size and in texture
of fur; forelimbs very small; above variegated with black, brown,
and yellow; pale yellow on sides and about eyes; belly grayish
white; forelimbs black on upper part. Head and body, 495 mm.;
tail, 330 mm. (Gould, 1841a, pp. 93-94.)
The former range included the interior of New South Wales and
Victoria, and the Murray River region of South Australia.
Gould writes (1863, vol. 2, p. 67) : "I have but little doubt that
this animal enjoys a wide range over the interior of New South
Wales; it certainly inhabits the Liverpool Plains as well as those
in the neighbourhood of the Namoi and the Gwydyr, from all of
which localities I have received numerous examples." He adds: "I
usually found it solitary, and sitting close in a well-formed seat
under the shelter of a tuft of grass on the open plains."
" According to Krefft, this species is common in the level country
between the Murray and Darling rivers" (Lydekker, 1894, p. 54).
E. Le G. Troughton (in litt., April 16, 1937) considers it "now
94
EXTINCT AND VANISHING MAMMALS
very rare, and apparently doomed to extinction in New South
Wales owing to denudation in over-stocked country, also populated
by rabbits, and the fox." In 1938 (p. 407) he refers to it as "either
extinct or nearly so."
In Victoria, according to C. W. Brazenor (in Hit., March 3, 1937) ,
there were two records, the last in 1869. He considers the species
extinct in that state.
"It is tolerably abundant in all the plains of South Australia,
particularly those situated between the Belts of the Murray and the
mountain ranges" (Gould, 1841J, text to pi. 12).
FIG. 13. Brown Hare-wallaby (Lagorchestes leporides). After Gould.
Jones writes (1924, pp. 222-223) of its status in South Australia:
"I know of no preserved specimens of this formerly common
animal from which a description may be written of the actual form
which inhabited this State. . . .
"In the British Museum catalogue of 1888 five specimens in the
collection are recorded as being from South Australia .... I have
been unable to obtain any evidence of its present existence in the
State, and in all probability it is completely exterminated."
Rufous Hare-wallaby; Western Hare-wallaby; "Whistler";
"Spinifex Rat"
LAGORCHESTES HIRSUTUS HIRSUTUS Gould
Lagorchestes hirsutus Gould, Proc. Zool. Soc. London 1844, p. 32, 1844.
("York District of Western Australia.")
FIG.: Gould, 1849, vol. 2, pi. 58.
This animal has long since disappeared from a large part of its
former range in the west of Western Australia but survives in
ORDER MARSUPIALIA: MARSUPIALS 95
indefinite numbers in the east along the South Australian border
and likewise in the northwestern part of the latter state.
General color of fur, especially on hind quarters and under parts,
rich sandy buff; head and back grizzled with grayish white; body
beset, especially posteriorly, with numerous long, rich rufous hairs;
space about eye reddish buff; ears large, grayish brown externally;
feet yellowish buff. Total length, 698 mm.; tail, 266 mm. (Gould,
1844a,p. 32.)
Stirling and Zietz (1893, pp. 154-155) record four specimens from
Western Australia ; they were taken during the Elder Expedition "in
the Porcupine grass (Triodia irritans) country, south of the Barrow
Range, before the exploring party entered the Victoria Desert. . . .
"Mr. Streich informs us that this animal appears to be numerous
in the northern parts of the Victoria Desert, where it often falls the
prey to the Wedge-tailed Eagle."
In an anthropological report on the same expedition, Helms says
(1896, pp. 240, 255-256) :
The Blyth Range, Barrow Range, and Victoria Desert tribes inhabit "spini-
fex country," where subsistence is difficult to maintain, and but for the
numerously occurring Largochestes [sic] hirsutus . . . and some other small
marsupials, it would probably be impossible for them to live in such desolate
districts. It can scarcely be wondered at that the majority of them appeared
lean and starvation-stricken. . . .
The Largochestes is almost, if not totally, absent here [in the vicinity of
Victoria Spring] ; and some 150 to 200 miles to the north, the eagle-nests,
which we had previously noticed daily, disappeared, which at once proved
the decrease of these small marsupials. . . .
Throughout the greater part of the interior, as far as the Expedition
went, their [the blacks'] principal flesh-food is supplied by the small marsupials
that harbor under the Zn'ocfoa-tussocks, and are commonly called "spinifex
wallabies" (Largochestes hirsutus), occurring abundantly in many places in
the triodia-regiou. . . . The blacks are very expert at killing the animals with
the "turtimbo," or short throwing-stick .... To enable them better to get
at this game they constantly burn large patches of the "spinifex" grass.
Shortridge (1910, p. 819; map, p. 820) considers the "mainland
form almost, if not entirely, extinct. Said possibly to still occur
very sparingly on sand-plains to the east of Beverley and York
where within quite recent times it was fairly plentiful.
"A single specimen was recorded from Hastings, near Kojonup,
in 1896, by the Perth Museum."
Glauert states (1933, p. 27) : "It has long disappeared from the
York district, where the first specimens were collected by Gilbert,
but survives in the desert country near the South Australian border,
and along the Canning Stock Route."
Finlayson (1935, pp. 63-67) gained experience with the species
in the northwest of South Australia. "The maala [L. hirsutus],
though common in the more westerly spinifex tracts, is not often
taken east of the [Aboriginal] Reserves." Much time was spent in
96 EXTINCT AND VANISHING MAMMALS
1932 in an unsuccessful search for it. "A year later, ... on the
south side of the Musgraves, we learned from the blacks that there
was a small colony of maalas in a spinifex patch ten miles south
of Koonapandi."
Finlayson then gives an account of hunting in this spinifex patch,
about 10 miles square, with some blacks. Their favorite method
of firing the country was utilized. As the fire drove the maalas out
of the tussocks, their chance of dodging the throwing-sticks was
slender. Those that escaped the fire by remaining in their burrows
were hunted out after the fire had passed. A satisfactory series of
specimens was secured.
The Rufous Hare-wallaby "requires observation and close pro-
tection wherever possible" (E. Le G. Troughton, in litt., April 16,
1937).
[It is represented by doubtfully distinct insular subspecies on
Dorre and Bernier Islands in Sharks Bay, Western Australia L. h.
dorreae Thomas and L. h. bernieri Thomas. Evidently both are
protected by their insular environment, and at least the latter is
reported as plentiful (Shortridge, 1910, p. 819).]
Banded Hare- wallaby; Banded Wallaby
LAGOSTROPHUS FASCIATUS (Peron and Lesueur)
Kangurus Fasciatus Peron and Lesueur, in Peron and Freycinet, Voyage Terres
Australes, vol. 1, p. 114, atlas, ed. 1, pi. 27, 1807. (Bernier Island, Sharks
Bay, Western Australia.)
FIGS.: Peron and Freycinet, Voyage Terres Australes, atlas, ed. 1, pi. 27,
1807, ed. 2, pi. 57, 1824; Gould, 1842, pi. 30; Waterhouse, 1846, vol. 1, pi. 4,
fig. 2; Gould, 1849, vol. 2, pi. 56.
Although Shortridge says (in Thomas, 1907, p. 772) that he had
never seen any animal, not even rabbits, in such numbers as this
species on Bernier Island, it now seems a melancholy necessity to
include it among the vanishing mammals of Australia. It is no
longer common on the islands in Sharks Bay where it once swarmed,
and it occurs in only a few isolated localities on the mainland of
Western Australia. (The mainland form, sometimes recognized as
L. /. albipilis (Gould) , is doubtfully distinct, and will not be treated
separately here.)
The animal is about the size of the Common Hare of Europe;
fur very long and soft, brown-gray, variegated with rusty, black,
and white; space about eye bright rusty; numerous narrow dark
cross bands on the back, most conspicuous posteriorly ; on the upper
parts and sides are very long interspersed white hairs; under parts
dirty white; hind feet with long, harsh, brownish-white hairs spring-
ORDER MARSUPIALIA: MARSUPIALS 97
ing from sides of two larger toes. Head and body, 444 mm.; tail,
279-305 mm. (Waterhouse, 1846, vol. 1, pp. 87, 90.)
At the time of its discovery by Peron in 1801, the species occurred
in great numbers on the islands in Sharks Bay (Bernier, Dirk Har-
tog's, and Dorre). A little more than a century later Shortridge
(in Thomas, 1907, p. 772) found the animals swarming on Bernier
Island. "It has been a particularly dry season, and they were very
thin. Food was evidently insufficient for them all, and dead speci-
mens were lying about in all directions. It would seem that they
have no natural enemies on the island; and they breed to such an
extent that the island will carry no more, and in times of drought
a number have to die." He adds (1910, p. 818) : "It may be noted
that sheep had been temporarily introduced there, while in the
south of Dirk Hartog there is a large sheep station, and the wallabies
are said to have entirely left that end of the island."
Glauert (1933, p. 27) reports the species as "not common" on the
islands in Sharks Bay.
On the mainland of Western Australia Gilbert found it in densely
thick scrubs, where "thie only possible means of obtaining it is by
having a number of natives to clear the spot, and two or three with
dogs and guns to watch for it. ... The natives are in the habit
of burning these thickets at intervals of three years, and by this
means destroy very great numbers." (Gould, 1863, vol. 2, p. 65.)
Thomas (1888, p. 182) recorded specimens from Wongar Hills,
York, and Perth.
Shortridge (1910, p. 818; map, p. 817) found it "existing in a
few isolated localities to the east of Pinjelly and Wagin, and accord-
ing to natives the Pellinup and Salt River districts in the neighbour-
hood of the Stirling Ranges.
"Plentiful enough in the restricted areas in which they occur,
frequenting thick prickly scrub."
He also remarks (pp. 818-819) on the "most sudden and unac-
countable" disappearance of this and a number of other mammals
in the Western, South-Eastern, and Central districts of Western
Australia; it "is said to have been first noticed about 1880." Short-
ridge continues:
The above areas are now, with a few exceptions, entirely devoid of indigenous
mammals. This is said partly to account for the way in which the natives have
been disappearing from the Western and Central districts of late years. . . .
The entire disappearance of so many species, over such large tracts of country,
is generally considered to be due to some epidemic or disease .... It may be
noted, however, that they have died out chiefly in the drier parts of the country,
where, except for the introduction of sheep, there has been very little altera-
tion in the natural conditions. Rabbits, although already very numerous in
the Centre and South-East, have not yet found their way to the North- West.
The mammals of the South- West, to about as far north as the Moore River,
... are rapidly retreating before civilisation. . . . The burning of forests
98 EXTINCT AND VANISHING MAMMALS
and general clearing of the country, together with constant raids of dogs
and domestic cats, are among the chief causes.
Glauert (1933, p. 27) records the mainland form as rare, occurring
"in a few isolated localities to the east of the Great Southern
Railway."
E. Le G. Troughton (in Hit., April 16, 1937) regards the extinction
of the mainland form as probable and states that suitable reserva-
tions must be made if the fauna is to survive on islands large enough
for commercial use.
Bridled Nail-tailed Wallaby
ONYCHOGALEA FRAENATA (Gould)
Macropus jraenatus Gould, Proc. Zool. Soc. London 1840, p. 92, 1841. ("In-
terior of New South Wales.")
FIGS.: Gould, 1841d, pi. 3; Waterhouse, 1846, vol. 1, pi. 4, fig. 1; Gould, 1849,
vol. 2, pi. 54; Le Souef, 1923, pi. 15; Australian Zoologist, vol. 3, pt. 4,
pi. 20, 1923; Le Souef and Burrell, 1926, fig. 48.
This beautiful wallaby, perhaps never very common, is threatened
with extinction in its ancient home in eastern Australia.
Form slender ; fur soft and short ; general color gray ; a white cheek
stripe ; sides of neck washed with cream ; a white stripe on each side
extending from occiput over the shoulder to join the white of the
under parts behind the arm insertion; space between these stripes
blackish brown ; tail black above and below toward tip, with a small
terminal nail; limbs whitish, darker on hands and feet. Total length,
825 (female) to 1,104 mm. (male) ; tail, 380 (female) to 482 (male).
(Gould, 1841a, p. 92, and 1841d, text to pi. 3.)
Its former range extended from southern Queensland to Victoria.
"0. fraenata inhabits the brigaloe-scrubs of the interior of New
South Wales and Queensland, and probably South Australia" (Gould,
1863, vol. 1, p. xxi). Gould (1863, vol. 2, p. 62) writes further of
its occurrence:
It is a native of the south-eastern portions of Australia, and the locality
nearest to the colony of New South Wales in which I observed it was Brezi,
on the river Mokai, whence it extended into the interior as far as I had an
opportunity of proceeding; Mr. Gilbert subsequently discovered that it was
common in the thick patches of scrub which are dispersed over all parts
of the Darling Downs. It inhabits all the low mountain ranges, the eleva-
tion of which varies from one to six hundred feet, and which are of a sterile
character hot, dry, stony, and thickly covered with shrub-like stunted
trees. . . .
In the neighbourhood of Brezi the natives hunt this species with dogs,
and often kill it with spears, bommerengs and other weapons; at Gundermein
on the Lower Namoi I found myself among a tribe of natives who succeed
in capturing them with nets ....
Its flesh, like that of the other small Kangaroos, is excellent, and when
procurable was eaten by me in preference to other meat.
ORDER MARSUPIALIA: MARSUPIALS
99
"There are probably only three species of animals that are entirely
confined to the fox area of Eastern Australia. These require our
immediate attention if the remnants are to be saved. They are the
Bridle Nail-tailed Kangaroo . . . , the Brush-tailed Rock-Wallaby
. . . , and Gaimard's Rat-Kangaroo. . . .
"The only Bridle Nail-tailed Kangaroos that exist as far as 1
know, are a few on Mr. Charles Baldwin's farm, near Manila, and
some in Taronga Park. Attempts to get this species to live in a wild
state in Taronga Park have failed, as they apparently cannot live
in the tick area, their proper home being the foot hills of the Dividing
Range of Eastern Australia." (Le Souef, 1923, p. 110.)
FIG. 14. Bridled Nail-tailed Wallaby (Onychogalea fraenata). After Gould
and photo by Berridge.
"Occasionally one . . . will be picked up by the great wedge-
tailed eagle. Remains of the animal have been found in the bird's
aerie. . . .
"This species, like so many other animals found in the more closely
settled parts of Eastern Australia, is now becoming very scarce, and
will probably soon be extinct. With the occupation of the land by
sheep and cattle, and the competition of the rabbit, the food and
shelter to which the wallabies were accustomed are decreasing. At
the same time, their enemies are increasing as their possible living-
grounds are becoming more and more restricted. But it is the
imported fox that is making the wholesale clearance, threatening
early extinction." (Le Souef and Burrell, 1926, pp. 210-211.)
"It is not uncommon in some parts of Southern Queensland, and
its pelts were frequently seen in the sales two or three years ago
100 EXTINCT AND VANISHING MAMMALS
under the name of 'padmelon.' It is now a protected species." (Long-
man, 1930, p. 59.)
Finlayson reports (1931, p. 85) on its status in the Dawson Valley,
Queensland: "Observed twice only, and no specimens obtained. It
was obtained by Lumholtz in the Rockhampton district in 1880-
1884, and recently Longman has stated that it is not uncommon in
South Queensland. Over the greater part of the Dawson country,
however, it is either absent or rare, as few reliable accounts of it
could be obtained."
"This gentle and beautiful species was once plentiful in inland
N. S. W. south to the Murray River, and in coastal parts as far as
Rockhampton in Queensland, but is now quite rare, or absent, over
entire range. A colony has been established on a small river island,
and such sanctuaries, free from foxes, probably represent the only
means of preventing extermination." (E. Le G. Troughton, in Hit.,
April 16, 1937.)
In Victoria there are a few records only, the last in 1867. The
animal is now extinct in that state. (C. W. Brazenor, in Hit., March
3, 1937.)
"As an illustration of the rapid breeding of marsupials, the ex-
perience of Mr. Chas. Baldwin, of Durham Court, Manilla, New
South Wales, is illuminating. In eighteen months Mr. Baldwin,
from five adults, bred seventy young of the bridled wallaby." (Hoy,
1923, p. 166.)
Crescent Nail-tailed Wallaby
ONYCHOGALEA LUNATA (Gould)
Macropus lunatus Gould, Proc. Zool. Soc. London 1840, p. 93, 1841. ("West
coast of Australia"; Thomas (1888, p. 78) lists the type specimen from
"Swan R., W. A.")
FIGS.: Gould, 1849, vol. 2, pi. 55; Lucas and Le Souef, 1909, p. 78, fig.
This wallaby is on the verge of extinction in the settled districts
of Western Australia but survives farther east toward the Great
Victoria Desert.
In general appearance it is very similar to the Bridled Nail-tailed
Wallaby but is slightly smaller; general color dark gray; face gray,
a mark over eye and cheek stripe slightly paler; a prominent white
crescent-shaped shoulder stripe, not encroaching on the neck; back
and sides of neck rich rufous ; a whitish hip stripe and another stripe
just above it; under parts whitish; tail uniform gray, its terminal
nail as in 0. fraenata. Male: head and body, 500 mm. ; tail, 332 mm.
(Thomas, 1888, pp. 77-78.)
The former range of this species included the southern parts of
Western Australia and Central and South Australia.
ORDER MARSUPIALIA I MARSUPIALS
101
"Mr. Gilbert's notes inform me that 'the Waurong ... is found
in the gum forests of the interior of Western Australia, where there
are patches of thick scrub and dense thickets . . . ; the dogs some-
times succeed in driving it out to the open spots, when, like the
Kangaroo rats, it runs to the nearest hollow log, and is then easily
captured' " (Gould, 1863, vol. 2, p. 64).
\\\f/*^^
FIG. 15. Crescent Nail-tailed Wallaby (Onychogalea lunata)
Shortridge (in Thomas, 1907, p. 768) considers it "very numerous
in some localities" of Western Australia. He adds (1910, pp. 815-
816, map) the following information:
"Within a more limited area this species seems to have much the
same range as Macropus eugenii, both forms frequently occurring
together . . . not extending far, if at all, beyond Beverley in the
North, or near the coast; its western boundary apparently being
the Darling Range.
"Also occurring in the southern interior of South Australia, where,
however, it is little known and probably rare."
Shortridge records 23 specimens from Arthur River and Woyaline
Wells.
102 EXTINCT AND VANISHING MAMMALS
"Mr. J. T. Tunney, whose fame as a collector is world-wide, in-
formed me that I could only hope to get the Crescent Wallaby . . .
along one obscure river [of Western Australia], and a forlorn hope
at that. Such a Wallaby should be energetically sought, trapped,
and placed, not in Zoological Gardens, but in the haven of a properly
supervised national reserve." (Troughton, 1923, p. 155.)
It occurs in "South-Western Australia, in isolated localities to the
west of the lower Great Southern Railway, probably on the verge of
extinction in the settled districts, but surviving further east towards
'the Great Victoria Desert" (Glauert, 1933, p. 29) .
It is "still found on the Nullarbor Plain" (A. S. Le Souef, in litt.,
February 15, 1937).
Jones (1924, p. 234) writes of its status as follows:
"In 1884 Mr. E. B. Sanger reported the Crescent-marked Wallaby
from the Centre and in the British Museum catalogue of 1888 three
South Australian specimens, collected by Sir George Grey, are
recorded.
"The Elder Expedition in 1891 met with it in the Everard Ranges
.... The Horn Expedition of 1894 obtained two specimens at
Alice Springs. I know of no more recent observations, and proba-
bly so far as South Australia is concerned the animal has ceased
to exist."
C. W. Brazenor writes (in litt., March 3, 1937) of a single Vic-
torian record, from the River Murray in 1857.
"Regarded as verging upon extinction in the settled areas of its
south Western Australian habitat, but surviving in the more desert-
like conditions between the Great Victoria Desert and Trans-Rail-
way, to the eastward. Extinction may be regarded as inevitable,
without establishment under favourable conditions." (E. Le G.
Troughton, in litt., April 16, 1937.)
Doubtless settlement and the concomitant imported pests have
accounted for the decline of this lovely wallaby.
Brush-tailed Rock-wallaby
PETROGALE PENCILLATA PENCILLATA (J. E. Gray)
Tufted-tailed or Mountain Kanguroo, K. pencillatiLs [J. E. Gray, in] Griffith,
Smith, and Pidgeon, Anim. Kingdom (Cuvier), vol. 3, Mammalia, pi.
opposite p. 49, 1827. (No type locality given.)
Kangwrus Pencillatus * [J. E. Gray, in] Griffith, Anim. Kingdom (Cuvier),
vol. 5, Mammalia, p. 204, 1827. ("New Holland" = "Sydney, N.S.W.,"
according to Iredale and Troughton, 1934, p. 42.)
FIGS.: Waterhouse, 1841, pi. 22, and 1846, vol. 1, pi. 1, fig. 1; Gould, 1842,
pi. 23, and 1853, pis. 39, 40; Lydekker, 1894, pi. 6; Le Souef and Burrell,
1926, fig. 47.
Formerly abundant in eastern Australia, this animal has suffered
serious reduction of range and numbers.
i Corrected to penicillatus in index volume, p. 23, 1835.
ORDER MARSUPIALIA: MARSUPIALS 103
The fur is long and thick; general color above dull brown, more
rufous on rump; an indistinct black mark behind the shoulder suc-
ceeded by a pale gray one; chin and chest pale gray; belly brown,
tinged with yellow; anal region yellowish rufous; arms and legs
rufous brown, becoming black at extremities; tail more or less
bushy, basal part rufous, remainder black, tip sometimes yellow.
Head and body, 720 mm.; tail, 560 mm. (Thomas, 1888, p. 67.)
The Brush-tailed Rock-wallaby once inhabited the ranges of
eastern Australia from southeastern Queensland to Victoria.
In New South Wales, according to Gould (1842, text to pi. 23),
"the species abounds wherever the kind of country suitable to its
habits occurs. The specimens in my own collection were procured
on the Liverpool range, and on the rocky sides of the mountains
facing Yarrundi on the Dartbrook, a tributary of the Hunter. I also
ascertained that it is very abundant on Turi, and the other moun-
tains situated to the eastward of the Liverpool Plains, and it doubt-
less ranges over a much greater extent of country than we are yet
acquainted with. It is ... strictly gregarious, assembling in such
numbers as to form well-beaten paths along the sides of the moun-
tains they inhabit. Their agility in leaping from rock to rock . . .
tends greatly to their protection, as neither the wily aborigine,
nor their still greater enemy the Dingo, can follow them to their
retreats . . . . "
Gould writes later (1863, vol. 2, pp. 46-47) :
"Those portions of the mountain ranges stretching along the
eastern coast from Port Philip to Moreton Bay . . . are among the
localities in which it is found; hills of a lower elevation than those
of the great ranges, and the precipitous stony gullies between the
mountains and the sea, are also situations it inhabits. . . .
"Of its flesh as an article of food I can speak most highly, having
frequently partaken of it in the bush and always found it excellent."
Le Souef calls attention (1923, p. 110) to the special need of
protective measures, since this species is entirely confined to the
fox area of eastern Australia. He adds that in New South Wales "a
few are found round Jenolan Caves, and at the head of the Murray
River." He also writes (1924, p. 272) : "The Brush-tailed Rock
Wallaby has become very scarce within Reynard's range during the
past few years. Skins of this species used to come into the sale
rooms in bales, now it is rare to see one."
Musgrave writes (1925, p. 210) of the species along the Nepean
River in New South Wales: "In former times the Brush-tailed Rock
Wallaby . . . occurred abundantly along the banks of the river, but
they have been so reduced in numbers by sportsmen that now they
are but rarely seen, and it is only a matter of time before the species
entirely disappears from the district."
104 EXTINCT AND VANISHING MAMMALS
"For all their agility . . . the rock-wallabies fall victims to very
sluggish enemies, for invariably the wallaby rocks are inhabited by
large carpet-snakes (Python varius) , which generally lie in wait
for their victims in the caves in which they take shelter" (Le Souef
and Burrell, 1926, p. 202).
Barry (1928, p. 163) reports a few in Kuringai Chase, near
Sydney, where, "despite protective laws, shooters, foxes and hounds
leave little chance of survival."
"The species should survive in the more rugged or inaccessible
parts of the Great Dividing Range in N. S. W., especially in some
of the sanctuaries recently declared, provided such are controlled,
and the public informed of dangers to survival of rarities, etc."
(E. Le G. Troughton, in litt., April 16, 1937) .
In Victoria it never occurred in great numbers and is now prob-
ably extinct. The last record was in 1905. (C. W. Brazenor, in litt.,
March 3, 1937.)
The species has been acclimatized on Kawau Island, New Zealand,
where it was introduced about 1870 and now exhibits some alteration
of coloration (Le Souef, 1930, p. 111).
[Petrogale herberti Thomas is treated as a subspecies of P. 'in-
ornata by Iredale and Troughton (1934, p. 43), who give its range
as "South Queensland (about 23 to 26 S. lat.)." However, it is
regarded as a subspecies of P. pencillata by Finlayson (1931, p. 82) ,
who writes of its status in the Dawson Valley: "Still . . . very
numerous and widely distributed. It is to be found in thriving
colonies in almost every range of hills away from the large towns."]
Yellow-footed Rock-wallaby; Bar-tailed Rock- wallaby
PETROGALE XANTHOPUS XANTHOPUS J. E. Gray
Petrogale xanthopus J. E. Gray, Proc. Zool. Soc. London 1854, p. 249, pi. 39
(Mammalia), 1855. ("Australia (Richmond River?)"; this is an erroneous
type locality, for Thomas (1888, p. 66) lists the cotypes from "Flinder's
Range, S. A.")
FIGS.: Proc. Zool. Soc. London 1854, pi. 39 (Mammalia); Gould, 1855, vol. 2,
pis. 43, 44; Royal Nat. Hist., vol. 3, p. 245, fig., 1894-95; Lucas and Le
Souef, 1909, p. 81, fig.; Le Souef and Burrell, 1926, fig. 46.
This largest and most striking of the Rock-wallabies has disap-
peared from many parts of its range in southern and eastern Aus-
tralia and is in urgent need of protection to prevent its extermination.
"Pale brown, minutely grizzled; chin and beneath white; streak
on side from back of shoulder, and along the side of the face under
the eye, whitish; dorsal streak narrow, brown; legs, feet, and tail
bright yellow; end of tail more bushy and varied with brown" (J. E.
Gray, 1855, p. 249). Fur long, soft, and silky; back of ears dark
ORDER MARSUPIALIA: MARSUPIALS 105
yellow; a brown blotch behind the elbow; a white patch on thigh
near knee; tail more or less annulated (Thomas, 1888, p. 65). Head
and body, 650-800 mm. ; tail, 600-650 mm. (Jones, 1924, p. 226) .
This animal's former range included southern and eastern South
Australia and the interior of New South Wales. It has also been
reported from Victoria and western Queensland.
Jones (1924, pp. 225-227) writes:
P. xanthopus inhabits the rocky country from the Gawler Ranges to the
Flinders Ranges, and to the eastern boundary of the State [South Australia]
at Bimbowrie and Cockburn. . . .
The Yellow-footed Rock Wallaby is still fairly abundant in certain parts
of South Australia, but from many of its old haunts it has completely dis-
appeared. It would seem that at the present time it is being driven mainly
to the north and east of the State, and that its last stronghold in South
Australia will be upon the New South Wales border. If it has not altogether
disappeared from the Gawler Ranges it must now be a very rare animal,
and in many parts of the Flinders Ranges its numbers are considerably
reduced. From the eastern portion of the State it will almost certainly
disappear before very many years are past, since its pelt is far too attractive
to permit it to survive as long as the fur trade exists. Although a totally
protected animal in this State, this protection is not extended to it by
certain of the States upon the borders of which it lives. It is therefore not
to be wondered at that pelts of the animal are disposed of in the markets of
States other than South Australia, even though the animal was obtained
within the geographical boundaries of our own State.
Petrogale xanthopus is a fitting example of an animal which needs sanc-
tuary for its preservation and more stringent legislative efforts to check its
slaughter.
Half a century ago Lydekker wrote (1894, p. 48) : "Some hun-
dreds of skins are annually imported to London from Adelaide,
their value ranging from one-and-fourpence each. The skins of the
common Rock- Wallaby [P. pencillata] are less valuable, averaging
from threepence to ninepence each, although they have been known
to reach as much as one-and-threepence."
"The [Melbourne] museum has no Victorian record of this species
though its range has been extended to the State in literature" (C. W.
Brazenor, in litt., March 3, 1937) .
A. S. Le Souef writes (in litt., February 15, 1937) that it is "very
scarce, probably nearing extinction owing to settlement and the fox."
"It provides an example of the need for unified control over State
protection, as pelts are sold in other States though the beautiful and
harmless marsupial is totally protected in South Australia. Such
action may cause extermination as the animal is being driven north-
east, and the hope of survival may rest with the sub-species de-
scribed from south-western Queensland." (E. Le G. Troughton, in
litt., April 16, 1937.)
[Petrogale celeris Le Souef, described from the vicinity of Ada-
vale, Bulloo River, southwestern Queensland, is regarded by Iredale
106 EXTINCT AND VANISHING MAMMALS
and Troughton (1934, p. 44) as a subspecies of P. xanthopus. Prac-
tically no information is at hand concerning its numerical status,
which, however, is presumably more satisfactory than that of P. x.
xanthopus.}
Red-necked Pademelon; Pademelon Wallaby
THYLOGALE THETIS (Lesson)
Hnlmaturus Thetis "Busseuil" Lesson, Manuel Mammalogie, p. 229, 1827.
("Port- Jackson" [Sidney], New South Wales.)
FIGS.: Geoffroy and Cuvier, Hist. Nat. Mamm., vol. 6, pi. 225, 1824; Lesson,
in Bougainville, Jour. Navigation Globe Thetis et Esperance, atlas, pi. 37,
1837; Gould, 1842, pi. 21; Gould, 1857, vol. 2, pis. 31, 32; Cabrera, 1919,
pi. 15, fig. 5; Le Souef and Burrell, 1926, fig. 42.
This species, formerly occurring from southern Queensland to
Victoria, is now extinct in Victoria, and its range in New South
Wales has become restricted to the north coast. Little seems to be
known of its present status in Queensland.
The upper lip is little developed, not hiding the front teeth. The
general color above is grizzled gray; rufous on neck, shoulders, and
rarely on cheeks and round .base of ears; a faint white hip-stripe
sometimes present; arms and legs gray or rufous; hands and feet
pale brown; tail gray basally, then brown above and white below.
Head and body, 540 (female) to 660 mm. (male). (Thomas, 1888,
p. 53.) Tail, 368 (female) to 476 mm. (male) (Waterhouse, 1846,
vol. 1, p. 148).
Of the early abundance of this pademelon, Gould writes (1863,
vol. 2, p. 38) :
Of the smaller species of Wallaby inhabiting New South Wales, the present
is perhaps the one best known to the colonists, inasmuch as it is more
abundant than any other. . . . All the brushes I have visited from Illawarra
to the Hunter, as well as those of the great range which stretches along
parallel With the coast, are equally favoured with its presence; I have also
received specimens from Moreton Bay. . . .
As an article of food, few animals are so valuable, its flesh being tender
and well-flavoured, and more like that of the common Hare than that of
any other European animal I can compare it with.
Le Souef and Burrell remark (1926, p. 196) that it "has been
noted in the scrub on the Blue Mountains, New South Wales."
Troughton states (1932, p. 188) that it shows "a continual shrink-
age in range" and is "now confined to the North Coast" of New
South Wales.
Lewis (1931, p. 120) apparently refers to the present species in
the following remarks on a Victorian animal: "The Dwarf Wallaby,
commonly known as the Paddymelon, was very plentiful once along
the coast between Lake Wellington and Metung, but I was afraid
ORDER MARSUPIALIA: MARSUPIALS 107
that these had been exterminated. I have ascertained, however,
that there are still some of them in this district."
In discussing the status of this and two other species in Victoria,
Jones and Manson say (1935, p. 35) : "All the small Wallabies are
rare and very infrequently seen."
More recently David H. Fleay (in Hit., June 1, 1937) reports
the species as extinct in Victoria but as still found in New South
Wales.
Some of the older works (e, g., Thomas, 1888, p. 53; Ogilby, 1892,
p. 53) record it from southern Queensland, and more recently Long-
man (1930, p. 58) lists it from southeastern Queensland. However,
Iredale and Troughton (1934, p. 46) give its current range as merely
"New South Wales."
Parma Wallaby or Pademelon; White-throated Wallaby or
Pademelon
THYLOGALE PARMA (J. E. Gray)
Hal [maturusl Parma "Gould" J. E. Gray, in Grey, Two Expeditions Discovery
Australia, vol. 2, appendix, p. 403, 1841. ("Sidney, and its neighbourhood,
New South Wales.")
Fia: Gould, 1856, vol. 2, pi. 28.
This species of New South Wales is evidently extinct.
The general color is deep reddish brown, penciled with white and
black; paler on sides; nape, shoulders, and forelegs brownish rust
color; a narrow black stripe along back of neck; throat and chest
white, rest of under parts dirty rusty white; tail scantily haired,
black above, dirty white below (Waterhouse, 1846, vol. 1, pp. 150-
151). Head and body, 590 (female) to 640 mm. (male) ; tail, 410
(female) to 430 mm. (male) (Thomas, 1888, p. 58).
Gould states (1863, vol. 2, p. 34) that in the Illawarra district
of New South Wales "I myself saw it in a state of nature. In these
extensive brushes it doubtless still exists, as since my return other
specimens have been sent to me from thence by the late Mr. Strange.
How far its range may extend westwardly towards Port Philip, or
eastwardly in the direction of Moreton Bay, I am unable to state."
Gould also speaks of its being hunted by the aborigines.
"This species seems to be very rare and locally distributed" in
New South Wales (Lydekker, 1894, p. 40).
"The White-throated Wallaby . . . once plentiful in the Illa-
warra district south of Sydney is apparently quite extinct, and rep-
resented by only five specimens of which three are in England and
two are in the Australian Museum." The uncertainty of range, as
expressed by Gould, "will never be cleared up now, as the last of the
two Australian Museum specimens was collected in 1889, and there
108
EXTINCT AND VANISHING MAMMALS
have been no recent evidences of its possible survival." (Troughton,
1932, p. 188.)
The range and status of the species are given by Iredale and
Troughton (1934, p. 46) as "New South Wales (south coast, possibly
extinct)."
A. S. Le Souef remarks (in Hit., February 15, 1937) : "I think
that this species is definitely extinct, though there may be a few
in the dense bush near Jervis Bay." He adds that recent search
and inquiry failed to reveal any trace of it.
FIG. 16. Parma Wallaby or Pademelon (Thylogale parma). After Gould.
Concerning the smaller wallabies in general, Le Souef and Burrell
say (1926, p. 195) : "The fur is fine and soft, and great numbers
are used for rugs, coats, and trimmings. We have practically no
knowledge as to the individual life-histories of this group."
Flinders Island Wallaby; Flinders Island Pademelon
THYLOGALE FLINDERSI Jones
Thylogale flindersi Jones, Mammals South Australia, pt. 2, p. 240, 1924 (cf.
Harper, 1940, p. 191). ("Flinders Island, . . . Investigator group, . . .
Great Australian Bight.")
This wallaby is confined to Flinders Island. "The colony was
estimated at a hundred or so in 1924, and in view of the presence
of rabbits as food destroyers, and cats, extinction seems certain
unless special measures are taken. This illustrates the need for
unsettled islands as sanctuaries, unless very large." (E. Le G.
Troughton, in Hit., April 16, 1937.)
The general color is grizzled light gray; sides and back of neck
and shoulders bright rufous in the male, tawny in the female; a
ORDER MARSUPIALIA : MARSUPIALS 109
well-marked pale area along the upper lip to beneath the eye; a
dark middorsal stripe from occiput backwards, very pronounced
in the male; chin and throat grayish white; lower neck, chest, and
belly colored almost like back, but somewhat lighter; limbs pale
fawn; tail pale gray. Head and body, 510 (female) to 570 mm.
(male) ; tail, 340 (female) to 410 mm. (male) . (Jones, 1924, p. 241.)
Jones gives the following account (1924, p. 242) :
Flinders observed this animal in 1802, and he records that on the island "a
small species of Kangaroo, not bigger than a cat, was rather numerous. I
shot five of them, and some others were killed by the botanists and their
attendants and found to be in tolerably good condition." Even comparatively
recently the animal was very numerous, and it has been reported that as
many as thirty thousand were killed on the island. In 1910 a destructive
bush fire swept the portion of the island occupied by a wallabies, and when
I visited the place in 1920 no traces of it were to be found, and the tenant
of the island believed it to be extinct. In 1922 I again visited the island and
found obvious evidences of its presence, but no actual specimen was seen.
In 1924 the little colony had considerably increased, and two specimens were
secured for study purposes. The present small colony of wallabies occupies
only a very limited area upon which the native bush has not been destroyed
by various attempts at cultivation. Although the colony probably contains
a hundred or so individuals its hold on life cannot be considered a very
secure one. It is always at the mercy of bush fires, having no line of retreat,
since it lives on a corner of the island that is girt by high and inaccessible
cliffs. Moreover, it has to contend against two introduced animals, the feral
domestic cat, which has overrun the island, and the food-destroying rabbit.
It may at any time, though fortunately this does not seem to be at present
the case, have to contend against human enemies. ... On account of its
build being rather more elegant than that of the thickset Kangaroo Island
wallaby it was at one time a favourite with people who cared to have
wallabies running in their grounds, but at present I believe there are no
descendants of these animals living on the mainland. It has also been an
inhabitant of the Zoological Gardens in Adelaide, but no specimens have
been exhibited there for many years. A former tenant of the island has
assured me that when the wallabies were numerous there were two distinct
types living in the island, the one obviously that described as Thylogale
flindersi, and the other a more rare, slender, yellow wallaby. What this
second species was it is impossible to guess; there seem to be no traces
of it left.
H. H. Finlayson writes (in litt., March 20, 1937) that although
the species is plentiful in a small area, its position is insecure.
Scrub Wallaby; Dam a Wallaby or Pademelon
THYLOGALE EUGENII (Desmarest)
Kangurus Eugenii Desmarest, Nouv. Diet. Hist. Nat., nouv. ed., vol. 17, p. 38,
1817. (Based upon the "kanguroo de File Eugene," Peron and Freycinet,
Voy. Terres Australes, vol. 2, p. 117, 1816; type locality "lie Eugene,
Josephine Archipelago," currently known as St. Peter's Island, Nuyt's
Archipelago, South Australia.)
FIGS.: Gould, 1841, pi. 11, and 1859, vol. 2, pis. 29, 30 (as Halmaturus der-
bianus); Lydekker, 1894, pi. 5; Le Souef and Burrell, 1926, fig. 43.
110 EXTINCT AND VANISHING MAMMALS
The present group of wallabies has long been in a state of great
taxonomic confusion, and part of the material necessary for elucida-
tion is evidently no longer obtainable. Under these circumstances
the group will be treated here as a specific unit, although attention
may be called to the three subspecies recognized by Iredale and
Troughton (1934, pp. 46-47) :
Thylogale eugenii eugenii (Desmarest). Type locality as stated
above.
Thylogale eugenii derbiana (J. E. Gray) . Type locality not stated
in the original description but said by Waterhouse (1846, vol. 1,
p. 155) to be "Swan River," Western Australia. Synonym: Macropus
gracilis Gould.
Thylogale eugenii binoe (Gould). Type locality: 'Tort Essing-
ton," Northern Australia. This is considered an error by Iredale and
Troughton (1934, p. 47), who substitute "Wallaby Island, Hout-
man's Abrolhos, West Australia" (cf, J. E. Gray, List of Specimens
Mammalia Brit. Mus., p. 91, 1843) ; however, Thomas (1888, p. 44)
lists the type from "Port Essington, N. T. (Sir J. Richardson) 1 '
and places binoe in the synonymy of Macropus agilis (Gould), as
Gould himself had already done (1863, vol. 2, p. 31). Synonyms:
Halmaturus houtmanni Gould; H. dama Gould; H. emiliae Water-
house.
The former range of the species as a whole included South Aus-
tralia, the coastal areas of southern and southwestern Western
Australia, and various islands along the coast, including Kangaroo
Island and Nuyt's Archipelago, South Australia, and the Recherche
Archipelago, Garden Island, and Houtman's Abrolhos, Western Aus-
tralia. It has become extinct on the South Australian mainland
and on St. Peter's Island (the type locality) ; in 1910 it was reported
as rapidly disappearing before settlement in Western Australia;
but apparently it remains plentiful in most of its insular habitats.
The following is adapted from Desmarest's description of what
may be considered the paratype, which presumably came from St.
Peter's Island: Fur soft; general color grayish brown, mixed with
rufous near the shoulders and on the nape, crown, and forelegs;
under parts whitish, distinctly separated from the dark color of the
upper parts; tail grayish brown above, white below, with a slight
reddish tint. Head and body about 21 (French) inches [567 mm.] ;
tail, a little more than 1 (French) foot [324 mm.].
Gould states (1841, text to pi. 11) that he had never heard of
"Halmaturus derbianus" being found on the mainland of South
Australia. But he writes (1863, vol. 2, p. 36) of its abundance on
Kangaroo Island. "The almost impenetrable scrub of dwarf Euca-
lypti, which covers nearly the whole of Kangaroo Island, will always
afford it a secure asylum, from which in all probability it will never
ORDER MARSUPIALIA: MARSUPIALS 111
be extirpated .... Such is the dense nature of the vegetation, that
nothing larger than a dog can follow it; still it is taken by men
residing on the island in the greatest abundance, both for the sake
of its skin and its flesh: they procure it principally by snares, a
simple noose placed on the outskirts of the brush; but they also
shoot it when it appears on the open glades at night."
Jones (1924, pp. 235-239) gives the following account for South
Australia :
Unfortunately the time has gone by when a good first-hand account of the
small scrub wallabies inhabiting South Australia could have been written.
The disappearance of the mainland wallabies is almost as remarkable a
phenomenon as the disappearance of the Native Cat. . . .
It is extremely difficult to define the former range of this complex species
on the mainland of South Australia, or even to discriminate with any cer-
tainty between the mainland form and the type of animal now living on
Kangaroo Island. Only a few years ago it swarmed in scrub-covered districts
all over the State, to-day it seems impossible to secure a single mainland
specimen for scientific study. In places where annual battues were held by
the present landowners less than twenty years ago it has disappeared altogether.
It is almost certain that some still linger upon the mainland, notably at the
southern end of Eyre's Peninsula and in the South-eastern districts, but so
far these animals have not been properly studied or preserved. . . .
In Kangaroo Island it is abundant and, since it lives in thousands upon
Flinders Chase fauna reserve, it is guaranteed, in so far as complete protec-
tion can guarantee it, perpetual survival. . . .
The wallaby of St. Peter Island has become extinct, and therefore we
cannot compare the animal now known as Thylogale eugenii with the St.
Peter Island animal, and, moreover, the original specimen described by
Desmarest is no longer in existence in Paris. It would seem to be somewhat
doubtful if the animal now known as Thylogale eugenii is the same as the
animal originally seen and captured on L'ile Eugene. The Kangaroo Island
Wallaby is readily kept and bred in confinement ....
E. Le G. Troughton remarks (in Hit., April 16, 1937) that its
survival "appears assured on Kangaroo Island, illustrating the value
of island sanctuaries in preserving remnants of vanishing stock."
Hoy writes (1923, pp. 164-165) of conditions on Eyre's Peninsula,
South Australia: "I was told, by a professional kangaroo hunter,
that at the time of the introduction of the fox he was always sure
of at least six dozen wallabies (Macropus eugenii) per week, but
during the season I was there, the fourth after the introduction of
the fox, he had not even seen one."
According to Finlayson (1927, p. 375), "the Thylogale of the
South Australian mainland has been exterminated before its identity
was properly established."
Shortridge (1910, pp. 812-813) gives its status in Western Aus-
tralia as follows:
"Very plentiful in many parts of the South- West, but rapidly dis-
appearing in the cultivated districts, especially towards the northern
end of its range. Not occurring in the coastal country between
112 EXTINCT AND VANISHING MAMMALS
Albany and Cape Leeuwin, although extending to the coast at the
Margaret River and Cape Naturaliste. Said still to exist in isolated
patches in the North between the Swan River and Gin-Gin. Also
occurring on the Abrolhos (Wallabi Group), Garden, and some
of the islands off Esperance. . . . Not extending on the South Coast
much beyond Phillips River . . . . "
Shortridge records specimens from Arthur River, Boyadine-Dale
River, Stockpool, Dwaladine, Woyaline Wells, Ellensbrook, and
Twin Peak and Middle Islands, off Esperance. The accompanying
map (p. 812) shows the former range extending in a broad coastwise
strip from Northampton to South Australia; but the current (1910)
range restricted to the southwestern corner of the state.
Under the name of Macropus (Thylogale) dama, Glauert (1933,
p. 32) gives the range of the mainland form as "South -Western
Australia, from the Moore River in the north to the south coast
(Cape Leeuwin and Cape Arid), inland to the Great Southern
Railway."
Troughton (1932a, p. 175) reports the species as "plentiful on the
two largest islands" of Houtman's Abrolhos.
Le Souef states (1930, p. Ill) that it was introduced about 1870
on Kawau Island, New Zealand, and is still present there.
Rufous-bellied Wallaby or Pademelon; Tasmanian Wallaby or
Pademelon
THYLOGALE BILLARDIERII (Desmarest)
Kangurus Billardierii Desmarest, Mammalogie, pt. 2, suppl., p. 512, 1822. ("La
terre de Van-Diemen" [Tasmania].)
FIGS.: Gould, 1841, pi. 10; Gould, I860, vol. 2, pis. 35, 36; Le Souef and Burrell,
1926, fig. 44.
Although this wallaby remains numerous in Tasmania and is still
found on some of the islands of Bass Strait, it seems desirable to
place it on record here as a vanished species of the Australian
mainland.
It is distinguished by its short ears, stout form, and long fur;
upper parts grayish brown, tinged with olive on head and rump;
under parts yellowish or rufous; tail short, grayish brown, the basal
part orange above, the terminal part grayish white below (Thomas,
1888, p. 59). Head and body, 640-765 mm.; tail, 315-320 mm. (Lord
and Scott, 1924, p. 247). Weight, 15-20 Ib. (Gould, 1863, vol. 2,
p. 42).
Gould gives the following account (1863, vol. 2, p. 42) :
I have but little doubt that the habitat of this Wallaby is limited to Van
Diemen's Land and the larger islands in Bass's Straits, in all which localities
it is so numerous that the thousands annually destroyed make no apparent
ORDER MARSUPIALIA: MARSUPIALS 113
diminution of its numbers. . . . Being one of the best-flavoured of the small
Kangaroos, it is very generally eaten in Van Diemen's Land.
The Tasmanian Wallaby may be regarded as strictly gregarious, hundreds
generally inhabiting the same localities .... It is very easily taken with snares,
formed of a noose placed in the run; and thousands are captured in this way,
solely for their skins : the sportsman also may readily procure it by stationing
himself in some open glade of limited extent, accompanied by two or three
small yelping dogs, before which it keeps hopping round and round, and thus
affords him an opportunity of shooting it as it passes.
B52L
Gunn states (1838, p. 106) that "they are excellent eating, but
the smallness of the skins renders them less valuable for tanning."
The recent status of the Rufous-bellied Wallaby in Tasmania is
given by Lord (1928, p. 19) : "It is evenly distributed and is plenti-
ful in certain districts remote from settlement; but close to the
settled areas its history is the same as the larger forms." He adds
(p. 23) that in recent years it has increased considerably in the
Tasmanian National Park. He also quotes (p. 24) the official
Tasmanian returns from the hunting of this species as follows:
1923 201,365
1924 86,393
1925 121,245
1926 94,531
Jones (1924, pp. 242-243) discusses its former occurrence in South
Australia :
In the collection of the British Museum there is a skull of this animal,
formerly the property of Sir Richard Owen, which came from Mount Gambier.
There are also two skeletons said to have been procured in South-east South
Australia. . . .
This is the common small wallaby of Tasmania, it is present also in some
of the islands of Bass's Straits and on the mainland of Victoria. Evidently
it was at one time an inhabitant of the South-eastern portion of this State,
where the Platypus and the Koala intruded into the South Australian fauna.
If it lingers in any corner of the South-East, I have been unable to ascertain.
I know of no South Australian specimens.
The species is now regarded as extinct in South Australia (David
H. Fleay, in litt., June 1, 1937).
Le Souef and Burrell, evidently referring to personal experience,
say (1926, p. 196) : "M. billardieri has been met with in Gippsland
(Victoria)."
C. W. Brazenor writes (in litt., March 3, 1937) : "Once a common
animal in southern Victoria, the species has now entirely disap-
peared, though it is still found on the islands of Bass Strait and in
Tasmania."
In the absence of the fox from Tasmania, this wallaby should
survive indefinitely in that country.
5
114 EXTINCT AND VANISHING MAMMALS .
Whiptail, Gray-face, or Pretty-face Wallaby; Parry's Wallaby
WALLABIA ELEGANS (Lambert)
Macropus elegans Lambert, Trans. Linnean Soc. London, vol. 8, p. 318, pi. 16,
1807. ("New South Wales.")
SYNONYM: Macropus parryi Bennett.
FIGS.: Lambert, op. cit., pi. 16; Bennett, 1835, pi. 37; Gould, 1842, pi. 19;
Gould, 1852, vol. 2, pis. 12, 13; Lydekker, 1894, pi. 4; Le Souef and Burrell,
1926, fig. 38.
This wallaby, well named elegans, is rapidly diminishing in num-
bers in its somewhat limited range in New South Wales and Queens-
land, and is in distinct need of total protection.
It is characterized by a slender and graceful build and a very
long tail. The general color is clear gray, with a bluish tinge; top
of muzzle brown, sides darker; white cheek-stripe sharply defined,
bordered below by a gray band; ears brown at base and tip, with
an intervening white area; digits of hands and feet black; under
parts grayish white; tail pale gray, with a black or gray crest below
the tip. (Thomas, 1888, p. 39.) Head and body, 732 (female) to
793 mm. (male) ; tail, 858 (female) to 1,077 mm. (male) (Finlayson,
19316, p. 77).
"With this animal neither the colonists of New South Wales nor
the naturalists of Europe are very familiar; not so much in con-
sequence of its being really scarce, as from the extreme shyness of
its disposition, the fleetness with which it escapes from its pursuers,
and the mountainous and almost inaccessible parts of the country it
inhabits. I did not succeed in procuring it myself while in Australia,
it being confined, as far as I could learn, to the range of hills which
stretch along parallel to the coast from Port Stephens [New South
Wales] to Moreton Bay [Queensland], a part of the country not
visited by me. Like most other members of its race, it is easily
tamed, readily becoming familiar and docile." (Gould, 1842, text to
pi. 19.)
"Mr. Strange informs me that it inhabits the rocky ranges of the
Clarence [New South Wales], occasionally descending into the more
open broken country, where it frequents the ledges of rocks at an
elevation of 2000 feet .... So fleet is this animal, that it is only
with the assistance of the finest dogs that there is any chance of
procuring examples ; it surpasses in fact every other animal in speed,
and when fairly on the swing no dog can catch it." (Gould, 1863,
vol. 2, p. 18.)
Finlayson (19316, pp. 75-77) gives the following valuable account
of the species in the Dawson Valley, Queensland:
This magnificent species still occurs in large numbers in suitable tracts all
over the valley, but in the northern part of the area is rapidly diminishing.
In 1884 it was obtained by Lumholtz near Rockhampton and on Coomooboo-
ORDER MARSUPIALIA: MARSUPIALS 115
laroo, for instance, but is now quite unknown in the vicinity of the first place
and on the second has become rare. . . .
Typical of the whiptail habitats are the beautiful undulating upland parks
of the broad-leaved ironbark (Eucalyptus siderophlora) .... As a character-
istic example of this type of country might be cited the Grevillea plateau,
where parryi [=eleaans] is still in very large numbers. . . .
It is distinctly social in habit, and very likely truly gregarious, though it
would take closer and more prolonged observation than I was able to give
to determine the point. It certainly camps in rather large parties, 12 or 15
being frequently seen lying up together, but in the late afternoon, when
feeding begins in earnest, there is a tendency, I believe, for the larger males
and females to go off in pairs. At Drumburle, where I watched it most, they
were so numerous, however, that towards evening whole hillsides were dotted
with the members of these disbanded camps, and it was impossible to make
out the existence of any natural grouping. Old males are always solitary,
as in many other species. . . .
Like so many mammals living in open country they are very curious, and
their curiosity has earned them a reputation for stupidity amongst trappers
and shooters. It is said by such, that in winter when large "mobs" congre-
gate on the sunny side of the ridges, a dozen may be shot down one by one
before the rest make up their minds to go, provided the shooter does not
move from his position.
Locally it is regarded as an extremely fast wallaby, but as it is not hunted
with dogs to any extent, it is difficult to get data for comparison with other
species. . . .
In considering the future of this wallaby in Queensland, there are sound
reasons for anxiety. It is true that it is still numerous over a large area, but
no one with any knowledge of the fate of open country species elsewhere
would maintain that it will long survive the present rate of slaughter in the
cattle country of the Dawson. Where man is concerned its instinct for
self-preservation is almost nil, and as its colouration and habits make it a
most conspicuous animal at any time, its destruction is almost a mechanical
matter. It is very probable that the scores of thousands of whiptails which
are killed every year in coastal Queensland, represent, not the natural increase,
as is assumed locally, but rather the natural drainage of the species from
large areas of relatively poor feeding grounds into smaller areas which are
more attractive to it and which will support a denser population. When the
country is settled these "fur pockets" act as natural traps, and destruction
which appears to be local actually affects a much wider area, indirectly.
It is this factor of natural concentration which is largely responsible for the
element of unexpected suddenness which often marks the extinction of mammal
species before advancing settlement.
M. parryi is one of the most beautiful of Australian mammals, and is one
of the very few species which can be easily and freely observed under natural
conditions. It is to be hoped that its value will be recognised while there
is still time.
E. Le G. Troughton writes (in Hit., April 16, 1937) that this
nearest eastern ally of the extinct Toolach was once plentiful in the
more open coastal country from north of Sydney to the Rockhamp-
ton district of Queensland. It is becoming rare owing to the natural
shrinkage of habitat with settlement and to destruction for "sport"
and profit. It is more beautiful and observable than most species
and requires total continued protection to ensure its survival.
116 EXTINCT AND VANISHING MAMMALS
Toolach; Toolache; Grey's Wallaby
WALLABIA GRBYI (Waterhouse)
Macropus (Halmaturus) Greyi Waterhouse, Nat. Hist. Mammalia, vol. 1,
p. 122, 1846. ("South Australia." Iredale and Troughton (1934, p. 50)
give, as a restricted type locality, the "Coorong, fide G. F. Angas.")
FIGS.: Gould, 1852, vol. 2, pis. 18, 19; Finlayson, 1927, pis. 16, 17.
This very beautiful wallaby, a former inhabitant of South Aus-
tralia, is apparently extinct in a wild state. "One or two specimens
in the Zoo at Adelaide are supposed to be the last living specimens
of this species" (A. S. Le Souef , in Hit., February 15, 1937) .
General color pale ashy brown, tinted with yellowish; under parts,
legs, and feet pale buff-yellow; toes black; head gray; a pale yellow
cheek-stripe, bordered above with blackish and below with brownish ;
back of neck and back of ears pale rufous; tip of ears black; tail
very pale gray, brown-white beneath, and with a terminal crest of
dirty yellowish hairs. Head and body, 761 mm.; tail, 660 mm.
(Waterhouse, 1846, vol. 1, pp. 122-124.) Additional characters given
by Jones (1924, pp. 244-245) are: a white patch above eye; back
with 10 to 12 dark gray bands; an ill-defined pale hip-bar. Head
and body, 810-840 mm. ; tail, 710-730 mm.
From the time of its discovery this species seems to have been
almost entirely confined to southeastern South Australia, chiefly
between the Murray River and Victoria. "Both species [Wallabia r.
rufogrisea and W. greyi] appeared to have crossed the Murray, but
the extent of their tenure of the river flats is difficult to estimate,
and from this north-western part of their range they were early
driven, or greatly reduced, by the rapid advance of closer settlement.
Their former presence in the lower part of the county of Sturt is
vouched for by many residents of that part of the country still
living." (Finlayson, 1927, p. 364.)
"The species was not exclusively confined to South Australia, but
occurred also through a small strip of Victorian territory contiguous
to the border" (Finlayson, 1927, p. 366).
"Mr. Strange informs me that he met with this animal 'between
Lake Albert and the Glenelg. The kind of country in which it is
found consists of large open plains intersected by extensive salt
lagoons and bordered by pine ridges. ... I never saw anything
so swift of foot as this species: it does not appear to hurry itself
until the dogs have got pretty close, when it bounds away like
an antelope, with first a short jump and then a long one, leaving the
dogs far behind it. ... I have had twenty runs in a day with four
swift dogs and not succeeded in getting one.' " (Gould, 1863, vol. 2,
p. 25.)
"Many people can remember the time when Toolaches swarmed
ORDER MARSUPIALIA: MARSUPIALS
117
in the neighbourhood of Kingston. Being by far the fleetest of all
the wallabies, its chase was at one time a very popular form of
sport, and its beautiful pelts have been marketed in very large
numbers in the salesrooms of Melbourne. ... It is not correct to
say that this very fine and distinctly South Australian wallaby is
extinct, for at the present moment five or six individuals still exist.
Any effort to preserve this remnant must be made immediately and
with vigour if it is to be of any service whatever." (Jones, 1924,
p. 245.)
FIG. 17. Toolach (Wallabia greyi). From photo.
Finlayson furnishes an extensive account (1927, pp. 367-369),
from which the following excerpts are taken:
The Toolach ... in all parts of its range showed a marked partiality for
grass country .... In the typical desert country of the counties of Russell
and Buccleuch, where grass flats are few and far between, it occurred but
sparsely, and here appeared to be comparatively solitary, but in the lower
south-east, where richer soils permit a far greater development of grasses,
its undoubted instinct towards gregariousness asserted itself, and when the
country was first settled it was here established in a series of isolated colonies
.... The groups . . . showed marked partiality for certain quite restricted
areas, from which they were only driven by persistent persecution, and to
which they returned again and again. . . .
A considerable weight of evidence inclines me to the belief that in point
of numbers M. greyi fell far short of the four other species of Macropus in
the district. Although human persecution and the occupation of its chosen
118 EXTINCT AND VANISHING MAMMALS
country early reduced its numbers and broke up and dispersed its larger
colonies, it was still far from uncommon even as late as 1910, and scattered
bands were still to be found in suitable localities. The chief of these were
along the edges of the long strip of grass country extending from a little
north of Millicent to the vicinity of Bull Island and Reedy Creek, and
known locally as Avenue Valley, on the Biscuit Flat between Robe and
Kingston, the Mosquito Plain between Naracoorte and Penola, and in the
country between Clay Wells and Conmurra, and probably also in the sand-
hill country of its northern district. Its rapid disappearance in the last
twenty years may be attributed with some confidence to the invasion and
enormous increase of the English fox, which has been proved without doubt
to take a heavy toll of the young, even of the large kangaroos, and indeed in
the almost unoccupied desert country where man has had little influence on
its destinies, it seems that the fox has been the sole factor in effecting its
extermination. Its chief natural enemies [sic] before the advent of the white
man and the fox seems to have been the wedge-tailed eagle (Uroaetus audax),
which, like the latter, chiefly attacked the young. These attacks were by no
means always successful, and were sometimes thwarted by the courage of the
females. . . .
I learn from another source that small boys in a certain district were in
the habit of periodically visiting the sites of eagles' nests to recover the
scalps from the remains of young toolaches to be found lying underneath;
this at a time when a bonus of sixpence was paid on all marsupial scalps.
By 1923 the species had become exceedingly rare. Isolated pairs were no
doubt scattered through the rougher stringy-bark country, but the sole
remnant of the Toolach population which continued living in country and
under circumstances which might be regarded as typical of that formerly
obtaining, was a small band of perhaps fourteen individuals, located on the
south end of Konetta sheep run, some twenty-six miles south-east of Robe.
Public attention was first called to the rapidly approaching extinction of
the Toolach by Professor Wood Jones, who repeatedly stressed the urgent
need for rigid protection of this group at Konetta. In May, 1923, as there
appeared little prospect for effective conservation in the south-east, an organ-
ised attempt was made on a considerable scale to capture living specimens
for transference to the sanctuary on Kangaroo Island. This, and a later
attempt in 1924, failed in their main objective, since as a result of overmuch
driving the four examples obtained were either dead or died shortly after
capture, but were not altogether fruitless, as much-needed Museum material
was thus acquired.
The subsequent history of the species consists of a resumption of the
exterminating process. Owing to the extensive publicity given to the two
expeditions noted above, local attention was focussed on the Toolach to a
degree hitherto unknown. Much of this attention was sympathetic to the
idea of conservation, but the realization of the great rarity of the wallaby
roused the cupidity of an unscrupulous few> and that survivors of the 1924
attempt have been wantonly killed for the sake of the pelt as a trophy,
is an assertion based on the admission of at least one of the slayers. The con-
stant hunting of foxes with dogs over the Toolach country has been made the
excuse for some of this killing, the plea being advanced that it is impossible
to prevent the dogs running anything and everything that is put up. ...
Interrogation usually elicits the fact that "nothing spoils a dog like checking
him." This peculiar solicitude for the dog's training has borne very heavily
on the Toolach and still bears very heavily on his cousin the brusher. Occa-
sionally, however, a better spirit prevails, and recently a Toolach doe was
promptly rescued from two kangaroo dogs which had seized her, and, in the
ORDER MARSUPIALIA: MARSUPIALS 119
patient care of Mr. J. Brown, of Robe, she has survived the rough handling
received. She may well represent the last of her race in this State, as a
careful and extended examination of the beat of the Konetta band by the
writer in February of this year failed to reveal any recent traces, either
in the shape of tracks or dejecta, and the opinion is expressed by the resi-
dent who knows the country best that the band has been entirely extirpated.
The species is very poorly represented in Museums, and enquiries recently
instituted in all the States indicate that there are six skins and seven skulls
in the public collections of Australia.
Mr. Finlayson writes (in litt., March 20, 1937) that the species
is nearly extinct.
"This beautiful species presents the most tragic, and probably
prophetic, history of all the kangaroos since white settlement" (E.
Le G. Troughton, in litt., April 16, 1937). "The sum total of the
isolated protective effort apparently is a doe, rescued from kangaroo
dogs, which by now may represent the sole survivor of the species"
(Troughton, 1938, p. 407).
Black-gloved Wallaby; Western Brush Wallaby
WALLABIA IRMA (Jourdan)
Halmaturus irma Jourdan, C. R. Acad. Sci. [Paris], vol. 5, p. 523, 1837. ("Les
bords de la riviere des Cygnes, sur les cotes de Leuwin (Australasie)"n:
Swan River, Western Australia.)
SYNONYM: Macropus (Halmaturus) manicatus Gould (1841).
FIGS.: Gould, 1841, pi. 9, and 1852, vol. 2, pis. 20, 21; Le Souef and Burrell,
1926, fig. 39.
Though apparently remaining common for the present in its re-
stricted range in Western Australia, this species "requires observa-
tion and close protection wherever possible" (E. Le G. Troughton,
in litt., April 16, 1937) .
Head gray above; cheeks and lips yellowish white; black spot
under chin; back of ears brown; inside of ears yellow, the terminal
third black; crown brown; chest, neck, sides, and outer surface of
limbs light tawny-yellow ; wrists and tarsi yellow ; digits brown and
black; tail with a dorsal and ventral crest, mostly gray, blackish
toward the end, and tipped with white hairs. Head and body, 720
mm.; tail, 630 mm. (Jourdan, 1837, p. 523.) According to Thomas
(1888, p. 41), the general color is dark bluish gray; back of ears,
crown, and digits black.
"To what extent this pretty animal ranges over Western Australia
has not been ascertained, but we know that it is very generally dif-
fused over every part of the colony of Swan River, wherever sterile
and scrubby districts interspersed with belts of dwarf Eucalypti
exist ....
"Mr. Gilbert informs us that it may be ranked among the fleetest
of its race; that it requires dogs of the highest breed to capture it,
120 EXTINCT AND VANISHING MAMMALS
and that a full-grown male weighs nearly twenty pounds. The flesh
forms an excellent viand for the table, and the skins manufactured
into rugs are extensively used by those whose avocations and mode
of life lead them to spend much of their time in the bush." (Gould,
1863, vol. 2, p. 27.)
Thomas (1888, pp. 41-42) lists specimens from Swan River,
Toodyay, and Perth.
Shortridge gives the following account (1910, pp. 809-811, map) :
Range almost identical with that of Macropus giganteus [= ocydromus],
except that it does not seem to occur in the southern coastal districts between
Cape Naturaliste and the Leeuwin. . . . Generally considered to be the
best sporting animal in Western Australia.
Not apparently dying out or disappearing even in the more thickly populated
districts to the same extent as the smaller marsupials.
Extending northwards beyond Watheroo, its range probably ends at some
point to the south of Geraldton.
Shortridge also records specimens from King River, Mount Barker,
Boyadine-Dale River, Stockpool, Dwaladine, and Woy aline Wells.
His map shows the range extending through the southwestern corner
of Western Australia between Geraldton and Esperance.
Le Souef and Burrell state (1926, p. 190): "The black-gloved
wallaby is still very numerous in South-west Australia. . . . This
species, strangely enough, is difficult to keep in captivity ; evidently
it requires special food to keep it in health." They also remark
(pp. 188-189) that "all wallabies live in or about scrub or brushwood,
for they have three enemies ever on the look-out for stray animals
that venture into the open, namely, the dingo, fox, and the great
wedge-tailed eagle."
According to Glauert (1933, p. 32) , the range of the present species
is "South-Western Australia, from the vicinity of Geraldton in the
north to the south coast. The eastern limit is approximately the
No. 3 Rabbit-proof Fence. Still common near Perth."
E. Le G. Troughton calls attention (in litt., April 16, 1937) to the
fact that this wallaby is the nearest relative of the practically extinct
Toolach, and suggests that "shrinkage of range should be watched
for, although survival seems at present assured by reservations, in
the event of more intensive cultivation."
Tasmanian Kangaroo; Forester Kangaroo; "Boomer"
MACROPUS TASMANIENSIS Le Souef
Macropus giganteus tasmaniensis Le Souef, Australian Zool., vol. 3, pt. 4,
p. 145, 1923. (Tasmania.)
Formerly common and ranging practically throughout Tasmania,
this fine species has been almost exterminated.
ORDER MARSUPIALIA: MARSUPIALS 121
Similar in size and appearance to M. giganteus, but with some-
what coarser fur and showing greater variation in color ; upper parts
sooty, sooty gray, rusty brown, or rusty red as a rule rusty brown,
with grayish under parts. Head and body, 1230-1400 mm.; tail,
970-1000 mm. (Le Souef, 19236, pp. 145, 147.) Size very large,
form slender and graceful; tail gray, with terminal portion black;
male about one-eighth larger than female. (Lord and Scott, 1924,
p. 244).
"In Van Diemen's Land, among other places, it resorts to the
bleak, wet, and frequently snow-capped summit of Mount Welling-
ton." In this country it "forms an object of chase, and like the
Deer and Fox in England, is hunted with hounds; and twice a week,
during the season, the Nimrods of this distant land may be seen,
mounted on their fleet steeds, crossing the ferry of the Derwent, at
Hobart Town, on their way to the hunting-ground, where they
seldom meet without 'finding'." (Gould, 1863, vol. 2, pp. 2-3.) The
same author gives (p. 4) an account of a hunt for an old male
"Boomer" that led the hounds a chase of 18 miles on land, and then
swam in the sea for more than 2 miles before he became exhausted
and was killed. He also (p. 2) quotes R. C. Gunn to the effect that
while the species may be found in numbers at certain places where
food is abundant, yet it is not as a general rule gregarious and does
not travel from place to place in flocks.
Lord (1928, p. 18) gives the following account:
The Forester Kangaroo formerly roamed over the greater part of Tasmania
where conditions were suitable. It frequents, as a rule, more open country
than M. ruficollis, and this fact, together with its larger size, is undoubtedly
responsible for its decline. At the present time this species is met with only
in a few localities in Tasmania. In some instances, the owners of large estates
have taken an interest in the animal, and it is owing to the protection thus
received that groups of this species exist to-day in certain places in the island.
In other parts where there are scattered mobs, such as in the extreme
North-East of Tasmania, the advance of settlement is having its effect,
for although the species is totally protected by law, the fact must be recog-
nised that in the more distant country districts it is a matter of extreme
difficulty to enforce the game laws.
Although very much reduced in numbers the Forester Kangaroo does not
appear to be in any immediate danger of extinction, particularly if the
landowners who have protected it in the past continue to recognise the
variety as one worthy of being retained. Again, the species will probably be
bred in local zoological gardens, and there is still the further possibility of this
and other species being bred on a large scale and made an item of great
economic importance to the State.
More recently R. Boswell writes (in litt., May 13, 1937) that
the species, though still wholly protected by law, has now been almost
exterminated through excessive hunting. There has been economic
exploitation of its hide and flesh.
122 EXTINCT AND VANISHING MAMMALS
Order INSECTIVORA: Insectivores
Family SORICIDAE: Shrews
This family is nearly cosmopolitan in distribution but is absent
in the Australian region (including New Guinea). There are about
25 genera and several hundred species and subspecies. They are
animals of generally small size and secretive habits. A single form
is considered extinct.
Christinas Island Musk-shrew
CROCIDURA FULIGINOSA TRICHURA Dobson
Crocidura juliginosa, var. trichura Dobson, in Thomas, Proc. Zool. Soc. Lon-
don 1888, p. 532, 1889. (Christmas Island, eastern Indian Ocean.)
This is described as a small shrew, with skull and teeth closely
like those of the species C. fuliginosa of the mainland (Assam,
Tenasserim, and Malay Peninsula) . Dobson differentiated it mainly
on the basis of a longer tail, beset with long fine hairs, but later
collections showed that the tail as recorded by Dobson for his speci-
men (80 mm.) was much longer than the average of 10 other speci-
mens for which C. W. Andrews (1900) gives measurements. The
color is not described but is doubtless, like that of the related form,
of a dark gray. According to Andrews, the well-haired tail is the
best character. Measurements: head and body, 65-82 mm.; tail,
63-75 mm.; hind foot, 13-17 mm.
The original specimen was brought back to the British Museum
by the surveying-ship Flying-fish under command of Captain
Maclear in 1886. Later, in 1897, a number of additional specimens
were secured by Andrews (1900), who lists measurements in his
Monograph of Christmas Island, and remarks: "This little animal is
extremely common all over the island, and at night its shrill squeak,
like the cry of a bat, can be heard on all sides. It lives in holes in
rocks and roots of trees, and seems to feed mainly on small beetles."
In 1908, Dr. Andrews again visited the island, to see what changes
had taken place with the establishment of a settlement, clearing,
and agriculture, since 1897. He found (1909, p. 102) that the shrew
"is probably also extinct, at least no specimen was either seen or
heard during my visit." He implies that this may have been due in
part to cats, which had been introduced and had become numerous.
However, cats would seem hardly sufficient to account for the ex-
termination of a shrew, which they will kill but seldom care to eat.
It may be that agricultural use of the land has reduced the numbers
of the shrew about the settlement at Flyingfish Cove; nevertheless a
careful search with modern collecting methods might still reveal the
animal's presence. On the other hand, if it is actually gone, one
ORDER PRIMATES: PRIMATES 123
may invoke some introduced disease, which seems, as Andrews de-
scribes, to have been the reason for the extinction of the two native
rats, Rattus nativitatis and R. macleari (q. v.).
G.M.A.
Order PRIMATES: Primates
Family LEMURIDAE: Lemurs
This family is restricted to Madagascar and the Comoro Islands.
Six genera and 26 species and subspecies are recognized. While some
forms remain common, others have become greatly reduced in num-
bers, and one is evidently extinct. Owing to a steady reduction in
the forested area of Madagascar and to a certain amount of perse-
cution by the natives, the lemurs are faced with a rather uncertain
future. Consequently, accounts of all the forms are provided in the
following pages.
Miller's Dwarf Lemur
MICROCEBUS MURINUS MURiNUS (J. F. Miller)
Lemur murinus J. F. Miller, Icones Anim. et Plant., pi. 13, 1777. (Madagascar.)
SYNONYMS: Prosimia minima Boddaert (1784) ; Lemur prehensilis Kerr (1792) ;
Lemur pusillus E. Geoffrey (1796) ; Galago madaffascariensis E. Geoffroy-
Saint-Hilaire (1812) ; Cheirogaleus minor E. Geoffroy-Saint-Hilaire (1812) ;
Microcebus rufus Wagner (1839); Myscebus palmarum Lesson (1840);
Microcebus myoxinus Peters (1852) ; Chirogalus gliroides Grandidier
(1868); Microcebus minor griseorufus Kollmann (1911).
FIGS.: P. Brown, New Illustrate. Zool., pi. 44, 1776; J. F. Miller, Icones
Anim. et Plant., pi. 13, 1777; G. Shaw, Cimelia Physica, pi. 13, 1796;
Audebert, 1800, Makis, pi. 8; E. Geoffroy-Saint-Hilaire, Ann. Mus. Hist.
Nat. [Paris], vol. 19, pi. 10, fig. 3; Peters, Reise Mossambique, Zool., I,
Saugethiere, pi. 3, 1852; Major, 1894, pi. 1, fig. 2; Milne Edwards, Grandi-
dier and Filhol, 1897, pi. 259, fig. 6; Kaudern, 1915, pi. 2, fig. 3.
The comparative abundance of this species is indicated by the
fact that the Mission Zoologique Franco-Anglo-Americaine of 1929-
31 secured 43 specimens (Delacour, 1932, p. 220) more than of
any other Madagascar lemur.
Size very small; head rounded; muzzle short and pointed; eyes
large and brilliant; ears large and naked (Forbes, 1894, vol. 1, p. 55) .
"Two phases, rufous brown or gray. The first has the head rusty
brown ; orbital ring and upper lip black ; stripe between eyes and on
nose, grayish white; upper parts of body rufous brown; dorsal line
indistinct ; sides of body and outer side of limbs mouse gray washed
with rufous brown ; entire under parts and inner side of limbs white
. . . ; tail rufous brown . . . ; hands and feet gray. The other phase
is mouse gray above, the back washed with rufous, a rufous spot
over each eye; outer side of limbs mouse gray; entire under parts
124 EXTINCT AND VANISHING MAMMALS
white; tail pale rufous." Total length, about 300 mm.; tail, 150 mm.
(Elliot, 1913, vol. 1, p. 104.)
Schwarz (1931, p. 403) gives the range as "the whole of S.E.,
E. [=S.], and W. Madagascar, as far north as the Betsiboka River
.... Exact limits in central Madagascar not known, but probably
only found in the plains. Not extending farther north than Ft.
Dauphin on the east coast." He mentions (pp. 402-403) specimens
from: Fort Dauphin; Ankazoabo, Bara; Ambolisatra and Itampolo
Be, north of Tulear; Tulear; and Morondava.
Sibree (1915, p. 243) refers to this as one of the most beautiful
and interesting of Madagascar lemurids. It "is remarkable also for
its large and very resplendent eyes, for the eye admits so much
light at dusk that quite an unusual brilliance is produced."
Three specimens were captured in 1932 in the Manampetsa Re-
serve in the southwest (Petit, 1935, p. 474) .
"At Tabiky [inland from Cape St. Vincent], the mouse lemur
was apparently very common and numbers were brought in alive
by natives. ... On November 2, 1929, fifteen specimens were
brought to me." Remains of a Microcebus were found in a pair of
goshawks (Astur henstii) taken near Tabiky. (Rand, 1935, p. 95.)
Smith's Dwarf Lemur
MICROCEBUS MURINUS SMITHII (J. E. Gray)
Cheirogaleus Smithii J. E. Gray, Ann. Mag. Nat. Hist., ser. 1, vol. 10, p. 257,
1842. ("Madagascar"; type locality restricted by Harper (1940, p. 192)
to "a few miles north of Fianarantsoa, central Betsileo.")
FIGS.: Forbes, 1894, vol. 1, pi. 6; Beddard, 1902, p. 544, fig. 260.
This lemur was reported as tolerably abundant by Shaw in 1879
(p. 135). Although scarcely any later information is at hand, its
nocturnal habits and its penchant for the tops of the highest trees
have perhaps safeguarded it from serious depletion.
Gray's type description (1842, p. 257) is as follows: "Pale brown;
streak up the nose and forehead, the chin and beneath paler; tail
redder." Schwarz (1931, p. 401) distinguishes this subspecies from
M. m. murinus as follows: "Tail not longer or shorter than head
and body. Colour above reddish brown; an indistinct dorsal band
sometimes present. Facial streak accompanied on both sides by a
distinct black stripe which extends as far as but hardly beyond
the eyes." Both body and tail are about 180 mm. in length in a
specimen from Majunga, north of the Bay of Bombetoka, which
Lorenz-Liburnau records (1898, p. 445) as M. myoxinus.
Schwarz (1931, p. 403) records specimens from: Vohemar, NE.
coast; Mananara, Bay of Antongil; Mahambo, north of Foulpointe,
NE. coast; Anabama Forest, Lake Alaotra; Ivohimanitra, Tanala;
ORDER PRIMATES: PRIMATES 125
Vinanitelo, SE. Betsileo; and north of Fianarantsoa, central Betsileo.
He states the range as follows: "The whole of eastern, northern,
and east-central Madagascar, including the plateau, as far south as
Ft. Dauphin. Also the north-west, down to the Bay of Bombetoka."
If, however, Schwarz is correct (p. 402) in recording murinus from
Fort Dauphin, the range of smithii can scarcely extend quite so
far south.
G. A. Shaw (1879, pp. 135-136) gives the following account:
They inhabit a belt of forest-land stretching from the eastern forest into
the heart of Betsileo, a few miles north of Fianarantsoa, where they are
tolerably abundant. They live on the tops of the highest trees, choosing
invariably the smallest branches ....
Their food consists of fruit and insects and most probably honey. I have
frequently seen them catching the flies that have entered their cage for
the honey; and I have supplied them with moths and butterflies, which they
have devoured with avidity.
They are extremely shy and wild. Although I have had between thirty and
forty caged at different times, I have never succeeded in taming one. . . .
I have had none breed in captivity.
Kaudern (1915, p. 74) records several specimens (as M. minor]
from Ste. Marie de Marovoay on the Betsiboka River in the north-
west, and one specimen (as M. smithii) from Fenerive on the east
coast.
G. M. Allen (1918, p. 516) records a specimen from Didy, south
of Lake Alaotra.
Coquerel's Dwarf Lemur
MICROCEBUS COQUERELI (Grandidier)
Cheirogalus Coquereli Grandidier, Rev. Mag. Zool., ser. 2, vol. 19, p. 85, 1867.
("Morondava," west coast of Madagascar.)
SYNONYM: Microcebus coquereli Schlegel and Pollen (1868).
FIGS.: Schlegel and Pollen, 1868, pi. 6; Milne Edwards, Grandidier and Filhol,
1897, pi. 259, fig. 4; Beddard, 1902, p. 544, fig. 261; Elliot, 1913, vol. 1,
pi. 4, upper fig. (facing p. 145).
To judge by the small number of specimens recorded, this is one
of the rarest lemurs of Madagascar.
It is a little smaller than Phaner furcifer; above dark gray, washed
with rufous ; tail dark rufous, except at the base, where it is colored
like the back; under parts yellowish gray. Head and body, 210 mm.;
tail, 340 mm. (Grandidier, 1867a, p. 85.)
Grandidier (1867a, p. 85) had seven of these animals in his pos-
session. They were nocturnal and lived on leaves and fruit.
Schlegel and Pollen state (1868, p. 13) that the species inhabits
the most impenetrable forests. They had only a single specimen,
secured in the forests of Congony, inland from the Bay of Passan-
dava, in northwestern Madagascar.
126 EXTINCT AND VANISHING MAMMALS
According to Elliot (1913, vol. 1, p. 107), the range extends on
the west coast from Cape St. Vincent to Helville, in the vicinity of
the Bay of Passandava.
Only five specimens are reported by Delacour (1932, p. 220) as
collected by the Mission Zoologique Franco-Anglo-Americaine of
1929-31.
Geoffrey's Fat-tailed Lemur
CHEIROQALEUS MEDIUS MEDIUS E. Geoffroy-Saint-Hilaire
Cheirogaleus medius [E.] Geoffroy-Saint-Hilaire, Ann. Mus. Hist. Nat. [Paris],
vol. 19, p. 172, 1812. (Type locality not stated; restricted by Schwarz
(1931, p. 405) to "Ft. Dauphin, S.E. Madagascar.")
SYNONYM: Opolemur thomasi Major (1894).
FIGS.: E. Geoffroy-Saint-Hilaire, Ann. Mus. Hist. Nat. [Paris], vol. 19, pi. 10,
fig. 2, 1812; Major, 1894, pi. 1, fig. 1.
Extremely little information is available concerning this lemur,
but it is evidently one of the less common of the Madagascar forms.
Head broad; snout short; upper parts gray, with a wash of rusty
brown, the tips of the hairs silvery; a whitish band extending from
between the eyes to the naked nose-pad; a whitish half-collar on
each side of the neck; orbital ring and ears brownish black; under
parts, inner side of limbs, and hands and feet yellowish white. Head
and body of female, 232 mm.; tail, 195 mm. (Major, 1894, p. 20;
type description of Opolemur thomasi.)
Major (1894, p. 20) records three specimens (as 0. thomasi) from
Fort Dauphin. "Nothing is known of the distribution . . . north of
Ft. Dauphin in eastern Madagascar" (Schwarz, 1931, p. 405).
The Mission Zoologique Franco-Anglo-Americaine of 1929-31 col-
lected nine specimens (Delacour, 1932, p. 220).
"These little lemurs are apparently entirely nocturnal .... At
Tabiky [inland from Cape St. Vincent], I found them in a gallery
forest through savannah and dry brush." (Rand, 1935, p. 95.)
Neither Delacour nor Rand gave the subspecific determination of
their specimens.
Samat's Fat-tailed Lemur
CHEIROGALEUS MEDIUS SAMATI (Grandidier)
Chiroffalus Samati Grandidier, Rev. Mag. Zool., ser. 2, vol. 20, p. 49, 1868.
("Flumen Tsidsibon in littore occidentali Madagascar insulae.")
FIG.: Proc. Zool. Soc. London 1872, pi. 70.
The few known specimens of this lemur come from a limited
section of the west coast of Madagascar (Tsidsobon River to Mo-
rondava) .
ORDER PRIMATES: PRIMATES 127
Fur of body and tail rather short; dark gray above, fulvous below;
tail fat, faded rufous; a white stripe from forehead to nose; orbital
ring black. Head and body, 190 mm.; tail, 170 mm. (Grandidier,
1868, p. 49.)
Major (1894, p. 18) and Schwarz (1931, p. 405) record specimens
from Morondava.
Milius's Mouse Lemur
CHEIROGALEUS MAJOR MAJOR E. Geoffroy-Saint-Hilaire
Cheirogaleus major [E.] Geoffroy-Saint-Hilaire, Ann. Mus. Hist. Nat. [Paris],
vol. 19, p. 172, 1812. (Type locality not stated; restricted by Schwarz
(1931, p. 406) to "Fort Dauphin, S.E. Madagascar.")
SYNONYMS: Lemur commersonii Wolf (1822); Cheirogaleus milii E. Geoffrey
(1828); Ch. typicus A. Smith (1833); Mioxicebus griseus Lesson (1840);
Chirogalus adipicaudatus Grandidier (1868).
FIGS.: E. Geoffroy, Ann. Mus. Hist. Nat. [Paris], vol. 19, pi. 10, fig. 1, 1812;
Geoffrey and Cuvier, Hist. Nat. Mamm., livr. 32, pi. 188, 1821; Proc.
Zool. Soc. London 1879, pi. 9 (ssp.?); Milne Edwards, Grandidier and
Filhol, 1897, pi. 259, fig. 5; Elliot, 1913, vol. 1, pi. 5 (ssp.?).
The meager information we have concerning this lemur is an
indication of its rarity.
Upper parts varying from brownish gray to ashy brown; under
parts and inner side of limbs yellowish or whitish; orbital ring
black; nose light gray; hands and feet dark brown; tail pale rufous
or ashy brown, sometimes with white tip. Total length, 580 mm.;
tail, 275 mm. (Elliot, 1913, vol. 1, pp. 93-94.) Ears naked for their
distal half; color more grayish than in Ch. m. crossleyi (Schwarz,
1931, p. 405).
The respective ranges of Ch. m. major and Ch. m. crossleyi are
none too clearly defined in the available literature. The former
seems to occur in the south and west of Madagascar; the latter, in
the center and the northeast.
Elliot's statement (1913, vol. 1, p. 93) of the range of the present
form is probably not altogether accurate: "Eastern coast of Mada-
gascar; Fort Dauphin to Tamatave; also in the lower wooded
regions of Betsileo Province; and on the west coast from Tullare
[=Tullear] to Pasandava, Central Madagascar."
G. A. Shaw (1879, pp. 134-135) records a specimen (subspecies
not determined) from the forests on the eastern side of Betsileo. "Its
food consists of fruits and possibly honey .... It appears to be a
very uncommon animal, ... as this is the only specimen I have
been able to obtain, although I kept a man in the forest for two
months seeking for one after I had obtained this one."
Major (1894, p. 22) records specimens from Morondava in the
southwest and from Tamatave and Ankay Forest in the northeast.
128 EXTINCT AND VANISHING MAMMALS
"Milius' Mouse-Lemur, though a rare species, is widely distributed
in Madagascar" (Forbes, 1894, p. 51).
Kaudern (1915, p. 74) records a specimen from Ste. Marie de
Marovoay on the Betsiboka River, northwestern Madagascar (where
the animal is said to be rather rare), and several specimens from
Andranolava, north central Madagascar. (Here again, the speci-
mens do not appear to have been determined subspecifically.)
Delacour (1932, p. 219) reports only six specimens collected by
the Mission Zoologique Franco-Anglo-Americaine of 1929-31.
Crossley's Mouse Lemur
CHEIROGALEUS MAJOR CROSSLEYI (Grandidier)
Chirogaliis crossleyi Grandidier, Rev. Mag. Zool., ser. 2, vol. 22, p. 49, 1870.
("Forets est d'Antsianak," Madagascar.)
SYNONYMS: Chirogale melanotis Major (1894); Chirogale sibreei Major
(1896).
FIGS.: Proc. Zool. Soc. London 1872, pi. 71, nearer fig.; Forbes, 1894, pi. 5.
This subspecies is apparently even rarer than Ch. m. major.
Upper parts rufous, especially on the head, under parts whitish;
head very large, rounded; orbital ring black; inner surface of ears
covered with dark brown hairs; tips of ears bordered with black;
tail short and well furred. Body, 200 mm.; tail, 120 mm. (Gran-
didier, 1870, p. 49.) Ears hairy inside and out, with hardly a naked
tip; fore parts of body strongly washed with brownish (Schwarz,
1931, p. 405). The tail of the type specimen of crossleyi was evi-
dently defective; the types of "melanotis" and "sibreei" have a
total length of 490-500 mm., and a tail length of 225-250 mm.
(Elliot, 1913, vol. 1, pp. 95-96).
This lemur seems to be scarcely known except from the type
specimens of crossleyi, "melanotis" and "sibreei" These are, re-
spectively, from the forests east of Antsianak; from Vohima on the
northeast coast; and from Ankeramadinika, one day's journey to
the east of Antananarivo. Thus the known range extends from east
central to northeastern Madagascar.
Hairy-eared Mouse Lemur; Tufted-eared Mouse Lemur
CHEIROGALEUS TRICHOTIS (Gunther)
Chirogaleus trichotis Gunther, Proc. Zool. Soc. London 1875, p. 78, 1875. (On
the "way from Tamantave to Murundava," Madagascar.)
FIGS.: Gunther, 18756, pi. 15, p. 79, figs. 1, 2; Milne Edwards, Grandidier and
Filhol, 1897, pi. 259, fig. 9.
Since the single specimen of the Hairy-eared Mouse Lemur was
described in 1875, no subsequent specimen has turned up, despite
the extensive collecting that has been carried on in Madagascar in
ORDER PRIMATES: PRIMATES 129
the meantime. It seems fairly necessary, therefore, to class it
among the extinct species.
"Brownish grey; lower parts grey, with the hairs white-tipped.
A triangular spot in front of the eye black; median line of the
snout and lips whitish. Hands and feet grey, with white-tipped
hairs. Ears very short, hidden in the fur. The lower part of the
concha and the space before the ear covered with a tuft of very
long hairs. Tail rather shorter than the body, covered with short
hairs . . , . All the nails pointed, claw-like." Head and body, 152
mm.; tail, 149 mm. (Gunther, 18756, pp. 78-79.) "This species
differs from all the members of this genus in the tufts of hair
standing out from the ears and sides of head, above the ears" (Elliot,
1913, vol. l,p. 97).
Elliot's authority for giving (p. 96) the range of the species as
the "forests of Antsianak" is not apparent. The exact type locality
is unknown, and no information is at hand concerning the route fol-
lowed by Crossley, the collector of the type, on his way from Ta-
matave on the east coast to Morondava on the west coast.
Fork-marked Lemur. Maki a fourche (Fr.)
PHANER FURCIFER (Blainville)
L[emur] furdfer Blainville, Osteogr., Mammif., Primates, Lemur, p. 35, 1839.
(Type locality not stated originally; "probably the region of the Bay of
Antongil, N.E. Madagascar" (Schwarz, 1931, p. 407).)
FIGS.: Blainville, Osteogr., Mammif., Primates, Atlas, Lemur, pi. 7, 1839;
Schlegel and Pollen, 1868, pi. 5; Milne Edwards, Grandidier and Filhol,
1897, pi. 259, fig. 3.
The Fork-marked Lemur was formerly abundant in Madagascar
and is still fairly common.
Upper parts reddish gray ; outer side of limbs dark rufous ; throat
pale rufous; chin and rest of under parts yellowish; a black stripe
from lower part of back to crown, where it bifurcates, one branch
ending over each eye; hands and feet dark brown; tail bushy, dark
reddish brown with black tip. Total length about 600 mm.; tail,
350 mm. (Elliot, 1913, vol. 1, p. 109.)
"This species has been recorded by Pollen and van Dam from
various localities on the N.W. coast, north of the Bay of Bombetoka
(Bay of Ampasindava, Jangoa River, Kongony River), but also
farther south at Morondava, on the W. coast. Found by M. J.
Audebert at Tassumbe/ N.E. coast." (Schwarz, 1931, p. 407.) He
also states that "there is no definite record of the occurrence of this
species considerably south of the Bay of Antongil." He mentions
having examined specimens from Vohemar, NE. coast, and An-
doany, NW. coast.
130 EXTINCT AND VANISHING MAMMALS
This pretty species is found in abundance in the forests of western
Madagascar, and it also appears to inhabit the eastern part. The
natives state that it is very fond of honey. It is nocturnal and its
chase is extremely difficult. (Schlegel and Pollen, 1868, pp. 9-10.)
Twelve specimens were collected by the Mission Zoologique
Franco-Anglo-Americaine of 1929-31 (Delacour, 1932, p. 220) .
"Near Tabiky [inland from Cape St. Vincent] , in November, 1929,
I found the squirrel lemur fairly common about my camp in a gallery
forest through savanna and low dry brush. Usually found in pairs,
. . . they sometimes moved about rapidly through the tops of the
tall trees, but more often were in the lower trees and bushes ....
"In the rain forest on Mt. d'Ambre [in the extreme north] this
lemur was fairly common, and noisy throughout the night, but it
kept to the tops of the forest trees." (Rand, 1935, p. 95.)
Broad-nosed Gentle Lemur
HAPALEMUR SIMUS J. E. Gray
Hapalemur simus J. E. Gray, Cat. Monkeys, Lemurs, and Fruit-eating Bats
Brit. Mus., p. 133, 1870. ("Madagascar.")
FIGS.: Gray, Proc. Zool. Soc. London 1870, pi. 52; Milne Edwards, Grandidier
and Filhol, 1896, pis. 122 A, 122 D (fig. 1), 122 E.
Comparatively few specimens of this species seem to be known,
and the information concerning it (other than anatomical) is very
meager.
<r Nose broad and truncated; . . . back iron-grey, with a rufous
tinge; the hairs black, with a subapical rufous band, and the lower
part lead-coloured; throat whitish; patch on rump at base of tail
yellowish" (J. E. Gray, 1870, p. 133). Forbes (1894, pp. 82-83)
adds the following details: ears short, covered with long hair;
sides of head, neck, and region round the eyes lighter than the
back; lower back, sides of body, and outer surface of limbs sooty
gray, with here and there a wash of rufous; tail, except at base,
sooty gray; under side of body and inner side of arms pale sooty
gray; no patch of spines on the arm above the wrist as in H. griseus.
Schwarz (1931, p. 407) gives a total length of 900 mm.
"It would appear . . . that the distribution . . . includes the
whole forested region of eastern Madagascar. It has not been
recorded up to now from the north-west." Single specimens from
Nandihizana, central Betsileo, S.E. Madagascar, and from Passum-
bee, N.E. coast, are mentioned. (Schwarz, 1931, pp. 407-408.)
G. A. Shaw writes (1879, pp. 133-134) of a live specimen that
"came from the higher-level forests on the eastern side of the
Betsileo, among the bamboos, on which it appears in a great mea-
sure to subsist. ... I have tempted it with very many different
ORDER PRIMATES: PRIMATES 131
kinds of berries and fruits growing in the forest; but it would not
touch any of them." It fed steadily and regularly upon grass.
G. M. Allen (1918, p. 516) records two specimens from near Am-
batondrazaka.
An indication of the rarity of this species is the fact that no speci-
mens were reported by the Mission Zoologique Franco-Anglo-Ameri-
caine of 1929-31.
Gray Lemur
HAPALEMUR GRISEUS GRISEUS (Link)
L[emur] griseus Link, Beytr. Naturg., vol. 1, pt. 2, p. 65, 1795. (Based upon
"Le petit Maki gris" of Buffon (Hist. Nat., suppl., vol. 7, p. 121, 1789);
type locality, Madagascar. )
SYNONYMS: Lemur griseus E. Geoffroy (1796); Lemur cinereus Desmarest
(1820); Hapalemur schlegeli Pocock (1917).
FIGS.: Buffon, Hist. Nat., suppl., vol. 7, pi. 34, 1789; Audebert, 1800, pi. 7;
Schlegel and Pollen, 1868, pi. 3; Milne Edwards, Grandidier and Filhol,
1896, pis. 122 B, 122 D (fig. 2), 122 F.
Extremely little new information concerning this subspecies has
come to light during the past 70 years. It must have become
quite rare.
Upper parts light olive-brown, brighter on top of the head; rest
of head gray; cheeks, throat, breast, and inner side of limbs ochra-
ceous-white; tail a little darker than the back. Total length, 24
inches; tail, 13^ inches. (Schlegel and Pollen, 1868, p. 7; Forbes,
1894, vol. 1, p. 81.) The general grayish green of this subspecies
is contrasted with the reddish green of H . g. olivaceus. The presence
of a wrist gland distinguishes both of these forms from H. simus.
(Schwarz, 1931, p. 408.)
Schwarz (1931, p. 408) gives the range of the present form as
follows: "The whole south and west, and the dry central plateau
as far east as Lake Alaotra ; it also goes north beyond the Betsiboka
River in the north-west." He records specimens from the following
localities: Lake Alaotra and Ambatondrazaka, central northeastern
Madagascar; District Ambalavo, in the southeast; and Tany Ma-
landi, in the northwest.
According to Schlegel and Pollen (1868, pp. 7-8), this lemur in
northwestern Madagascar inhabits by preference the forests of
bamboo. It was found at a few days' journey from the coast, along
the Ambassuana River in the Tanimalandy district. It is entirely
nocturnal, and sleeps during the day on the highest stems of the
bamboos. The stomachs of all specimens were found filled with
bamboo leaves.
Delacour (1932, p. 219) records 25 specimens of "Hapalemur
griseus" as collected by the Mission Zoologique Franco-Anglo-Ameri-
132 EXTINCT AND VANISHING MAMMALS
caine of 1929-31. However, the further account given by Rand
(1935, p. 95) indicates that these represent the subspecies olivaceus.
Olivaceous Lemur
HAPALEMUR GRISEUS OLIVACEUS I. Geoffrey
Hapalemur olivaceus I. Geoffroy, Cat. Method. Mamm. [Mus. Paris], pt. 1,
Primates, p. 75, 1851. ("Madagascar." Type locality restricted by Elliot
(1913, vol. 1, p. 127) to "Ampazenambe, Madagascar.")
FIGS.: Forbes, 1894, vol. 1, pi. 8 (ssp.?) ; Milne Edwards, Grandidier and
Filhol, 1896, pis. 122 C, 122 D (fig. 3).
This subspecies appears to be considerably commoner than E.g.
griseus.
It is similar to the latter but has a longer and denser pelage;
color olive, with a rufous tint; throat gray rather than white;
cheeks speckled gray (I. Geoffroy, 1851, p. 75).
According to Schwarz (1931, pp. 408-409) , this lemur "is found in
the moist and wooded north-east and east [of Madagascar], prob-
ably as far south as Fort Dauphin." He records specimens from
the following localities: Mananare, Bay of Antongil; Vohemar,
NE. coast; Tamatave, E. coast; Analamazaotra, east of Tananarive;
Ambohimitombo, Tanala country, E. Madagascar; Vinanitelo, S.
Betsileo ; and several localities on the east coast north of Tamatave.
Kaudern (1915, pp. 70-71) reports the animal as apparently not
rare in the forests west of Fenerive on the east coast, where he
obtained three specimens from the natives.
Rand writes (1935, p. 95) that it is diurnal. He found it in
the tops of the lower trees in the rain forest, and occasionally in
dense thickets of bamboo on the edge of the forest. "Occasionally
found singly, it was more often seen in groups of two or three. At
Manombe in the southeast I saw two running about through the
forest tree tops .... Hapalemur was fairly common about camp
two days northeast of Maroantsetra."
Ring-tailed Lemur
LEMUR CATTA Linnaeus
[Lemur] Catta Linnaeus, Syst. Nat., ed. 10, vol. 1, p. 30, 1758. (Based upon
the "Maucauco" of Edwards (Nat. Hist. Birds, pt. 4, p. 197, 1751);
type locality, "Madagascar.")
FIGS.: Edwards, 1751, pi. 197; Schreber, Saugthiere, vol. 1, pi. 41, 1774; Aude-
bert, Hist. Nat. Makis, pi. 4, 1800; Geoffroy and Cuvier, Hist. Nat. Mamm.,
livr. 5, pi. 27, 1819; Milne Edwards and Grandidier, 1890, pis. 171, 172;
Royal Nat. Hist., vol. 1, p. 211, fig., 1893-94; Proc. Zool. Soc. London
1906, p. 124, fig. 48; Elliot, 1907, p. 545, fig. 76; Elliot, 1913, vol. 1, pi. 5,
upper fig.
ORDER PRIMATES: PRIMATES 133
This is perhaps the best known of all the Madagascar lemurs,
and it still appears to be moderately common.
The fur is soft and delicate; face and ears white; nose and area
about each eye black; top and back of head dark ashy; back and
sides of a redder ash color ; outer side of limbs light ashy ; upper sides
of paws whitish ; under parts and inner sides of limbs white ; tail with
broad alternate rings of black and white (Edwards, 1751, p. 197).
"Length of body and tail together, 40 inches" (Forbes, 1894, vol. 1,
p. 76).
"This species, which inhabits rocky open country, is found in
south-western, southern, arid south-eastern Madagascar" (Schwarz,
1931, p. 410).
"As far as my experience of seven years goes, these Lemurs are
found only in the south and south-western borders of the Betsileo
province of Madagascar." They are not found in the forests, but
among the rocks. "The prickly pear . . . constitutes their chief
article of winter food .... Their summer food consists of different
kinds of wild figs and bananas." (G. A. Shaw, 1879, pp. 132-133.)
This species bears captivity well, and is everywhere offered for
sale by the natives (Kaudern, 1915, p. 50).
Schwarz (1931, p. 410) records specimens from Tulear in the
southwest, and adds: "It has been found by van Dam at Moron-
dava, Matseroka, and the Bay of St. Augustin in the south-west,
and is recorded by him as far north-east as the region of Ft.
Dauphin."
This species ranges over a vast area, from Mangoky on the
north to beyond Menarandra on the south. Decary records it in
Androy along all the rivers and as far as the region of Beloha.
Perrier de la Bathie has observed bands on the western parts of tho
massif of Andringitra. (Petit, 1931, p. 560.)
Thirty specimens were collected by the Mission Zoologique Franco-
Anglo-Americaine of 1929-31 (Delacour, 1932, p. 219) .
Rand (1935, p. 96) mentions observations on this lemur at
Ampotaka and Lake Tsimanampetsotsa and near Tulear. He also
writes (pp. 95-97) : "The ring-tailed lemur was found in and about
most of the more densely -wooded areas and the gallery forest in the
arid parts of southwestern Madagascar.
"It is a diurnal and crepuscular creature ....
"This animal, like most of the lemurs, is gregarious. It was usually
seen in parties of from four or five up to ten or fifteen and more. . . .
"Wild specimens were seen to eat leaves of certain trees ....
"They are often kept as pets by Europeans and are carried to
various parts of the island."
At the Manampetsa Reserve in the southwest numerous bands
were seen in 1926, but the species was rare in 1933 (Petit, 1935,
p. 474).
134 EXTINCT AND VANISHING MAMMALS
Black Lemur
LEMUR MACACO MACACO Linnaeus
[Lemur] Macaco Linnaeus, Syst. Nat., ed. 12, vol. 1, p. 44, 1766. ("Mada-
gascar.")
SYNONYM: Lemur leucomystax Bartlett (1863).
FIGS.: Schreber, Saugthiere, vol. 1, pi. 40 A, 1774; Proc. Zool. Soc. London
1862, pi. 41; Schlegel and Pollen, 1868, pi. 1; Proc. Zool. Soc. London 1880,
p. 451, fig. 1; Proc. Zool. Soc. London 1885, p. 672, fig.; Milne Edwards and
Grandidier, 1890, pis. 130-132.
The Black Lemur still occurs in fairly large bands in the north-
west of Madagascar.
It has the ears tufted, with long hairs continuing down the side
of the neck to the angle of the mouth. The male is entirely black.
Female: snout and back of head black; forehead blackish gray;
whiskers and ear-tufts white; general body color rich ferruginous
brown; limbs and neck reddish yellow; tail whiter; under parts
and inner side of limbs creamy white. There is a considerable degree
of variation in the color of this species. (Forbes, 1894, vol. 1, p. 70.)
Total length, 41 inches; tail, 22 inches (Schlegel, 1876, p. 303).
"The range ... is limited to the forests of the N.W. coast, north
of the Bay of Bombetoka and the coast islands. It has been recorded
by Pollen and van Dam from the following places: Anorontsanga ;
Syrangene; Kongony and Jangoa Rivers; Andoany, Narendry Bay
("Maroandiana")." (Schwarz, 1931, p. 417.)
These animals inhabit the forests extending between the Bay of
Diego-Juarez and the Bay of Bombetoka, as well as the forest of
Loucoube in the isle of Nossi-Be. They live in bands in the highest
trees of the impenetrable forests. Bananas are their ordinary food
in the wild state. They are also fond of the brains of birds. (Schlegel
and Pollen, 1868, p. 2.)
The range includes the upper Sambirano (Petit, 1931, p. 562).
Eight specimens were collected by the Mission Zoologique Franco-
Anglo-Americaine of 1929-31 (Delacour, 1932, p. 220) .
"The parties observed were much larger than those of that species
[L. julvus], containing sometimes as many as 18 individuals
males, females, old, and young. They were always very tame."
(Rand, 1935, p. 99.)
Sanford's Lemur
LEMUR MACACO SANFORDI Archbold
Lemur julvus sanjordi Archbold, Am. Mus. Novit., no. 518, p. 1, 1932. ("Mt.
D'Ambre, Madagascar.")
This lemur is known only from 18 specimens collected in the
type locality by the Mission Zoologique Franco-Anglo-Americaine
of 1929-31.
ORDER PRIMATES: PRIMATES 135
It differs from all the black-nosed lemurs, except Lemur macaco
rujus, in its lighter color, and from the latter in its cheek and ear
tufts, in which it resembles Lemur macaco. Nose black; top of head
dark olive-buff; general color of upper parts snuff-brown to drab;
hands russet ; spot at root of tail and basal half of tail bay-colored ;
distal half of tail bushy, the hairs with blackish brown tips and with
a subterminal clay-colored band; a ruff of long hairs around the
ears and down the cheeks, varying from white to light ochraceous-
buff; under parts gull-gray, washed with buffy. Female without
cheek-tufts. Total length of male, 895 mm.; tail, 495 mm.
This form is restricted to the rain forest of Mount D'Ambre,
northern Madagascar. (Archbold, 1932, p. 1.)
White-fronted Lemur; White-faced Lemur. Maki a front
blanc (Fr.)
LEMUR MACACO ALBIFRONS E. Geoffrey
L[emur] Albifrons [E.] Geoffroy, Mag. Encycl. [2d yr.], vol. 1, p. 48, 1796.
(Type locality not stated in original description, but later given as "Mada-
gascar" (E. Geoffroy-Saint-Hilaire, 1812, p. 160).)
SYNONYM: Prosimia frederici Lesson (1840).
FIGS.: Audebert, Hist. Nat. Makis, pi. 3 (facing p. 13), 1800; Geoffroy and
Cuvier, Hist. Nat. Mamm., livr. 3, pis. 17, 18, 1819; Milne Edwards and
Grandidier, 1890, pis. 136, 144, 154, 155.
To judge by the 36 specimens collected by the Mission Zoologique
Franco-Anglo-Americaine of 1929-31 (Delacour, 1932, p. 220) , this
must be one of the commonest of Madagascar lemurs.
Pelage brown; top of head, cheeks, and jaws white (. Geoffroy,
1796, p. 48). "Of this race there are two mutations which occur
together, viz., (1) a normally black-headed, whitish-cheeked type,
with dark ground-colour, in which the female is only slightly paler
than the male, and (2) the 'albifrons' type, which shows a reduction
of black pigment, the ground-colour being more reddish, especially
so in the female; the male has the whole crown, cheeks, and beard
white or whitish, but there are females which show a whitening on
the head, although the normal phase of the 'albifrons' female has a
lead-grey head and a grey muzzle" (Schwarz, 1931, p. 410).
"The range of this local race apparently includes the north-
eastern coast of Madagascar as far as, and slightly beyond, the
Bay of Antongil. The northern and western limit is uncertain;
only the dark phase apparently occurs on the north-west coast."
Specimens are recorded from: Vohemar and Sahambavany, N.E.
coast; Mananare, Maroansetra, Androutse, and Ampazenardo, in
the vicinity of the Bay of Antongil. (Schwarz, 1931, pp. 410-411.)
"The effects of hunting by natives on the animal life of Mada-
gascar are negligible. The main destruction of the fauna is caused
136 EXTINCT AND VANISHING MAMMALS
by the cutting of the forests. But near Maroantsetra we saw what
might happen. Near the town itself these lemurs were scarce and
very wary, apparently having been hunted with guns, but once
away from large settlements they were very common and tame. On
July 22, 1930, two days northwest of Maroantsetra, I surprised a
party of eight in the low bushes of the ground cover in the forest.
They fled but a little way and I sat down to watch them. Very soon
the whole party came back and resumed feeding on the fruit of a
low bush near me.
"Parties of these lemurs were often heard grunting and growling
in the forest as though fighting." (Rand, 1935, p. 98.)
Brown Lemur; Fulvous Lemur. Maki brun (Fr.)
LEMUR MACACO FULVUS E. Geoffrey
L[emur] Fulvus [E.] Geoffroy, Mag. Encycl. [2d yr.], vol. 1, p. 47, 1796.
("Madagascar." This subspecies is considered typified by specimens from
the Tamatave region (Schwarz, 1931, p. 411).)
SYNONYMS: Prosimia macromongoz Lesson (1840); Lemur bruneus van der
Hoeven (1844).
FIGS.: Buffon, Hist. Nat., suppl., vol. 7, pi. 33, 1789; Lacepede and Cuvier,
1801, unnumbered pi.
Although it is said that "this race is about the most common
Lemur in captivity" (Schwarz, 1931, p. 412), very little information
can be offered concerning it, owing partly to the confusion that has
long prevailed in the taxonomy and nomenclature of this specific
group.
The pelage is brown above, gray below; head black; hands fulvous
or brown (]5. Geoffroy, 1796, p. 47). The animal is a third larger
than Lemur mongoz; its tail, less bushy and more woolly, tapers
toward the tip; rump and legs washed with olive (fi. Geoffroy, in
Lacepede and Cuvier, 1801, p. 3 of "Le Maki Mococo et le Maki
brun"). Ground color olive-brown; cheeks yellowish white (Schwarz,
1931, p. 410).
"The range of this race is not completely known. It obviously
inhabits the coast between the Bay of Antongil and Andovoranto,
but may go farther south, as far as Mahanoro. In the interior it
appears to go into the forest-belt east of Tananarive." Specimens
are recorded from: Andragoloaka, S.E. of Tananarive, Prov. Ime-
rina; Lakato Forest, Ankay, N.E. of Tananarive, Imerina; Sakana
and Ambotorao, opposite the He Ste. Marie; and Tamatave.
(Schwarz, 1931, p. 411.)
ORDER PRIMATES: PRIMATES 137
Collared Lemur. Maki a f raise (Fr.)
LEMUR MACACO OOLLARIS E. Geoffrey-Sain t-Hilaire
Lemur collaris [E.] Geoffroy-Saint-Hilaire, Ann. Mus. Hist. Nat. [Paris],
vol. 19, p. 161, 1812. ("Madagascar.")
SYNONYMS: Prosimia melanocephala J. E. Gray (1863); P. xanthomystax
J. E. Gray (1863); ?F. flavifrons J. E. Gray (1867); Lemur nigerrimus
P. L. Sclater (1880); Lemur mongoz var. cinereiceps Milne Edwards and
Grandidier (1890).
FIGS.: Geoffroy and Cuvier, Hist. Nat. Mamm., livr. 2, pi. 11, 1819; Proc.
Zool. Soc. London 1863, pi. 17 (as Prosimia xanthomystax), pi. 18 (as P.
melanocephala) ; ?Proc. Zool. Soc. London 1867, pi. 31 (as P. flavifrons) ;
?Proc. Zool. Soc. London 1880, p. 451, fig. 2 (as Lemur nigerrimus) ;
Milne Edwards and Grandidier, 1890, pis. 140, 147 (as L. mongoz var.
cinereiceps); Elliot, 1913, vol. 1, pi. 6 (as L. julvus).
This lemur appears to be common in southeastern Madagascar.
Pelage rufous-brown above, fulvous below; a ruff of rufous
hairs; face lead-colored (E. Geoffroy-Saint-Hilaire, 1812, p. 161).
Cheeks orange-yellow; ground color light brown; a faint spinal
line generally present; female generally with head gray. "This
race . . . may be slightly smaller than the other races .... There
is considerable individual variation, especially as regards the amount
of black, grey, or reddish brown on the crown and forehead. There
are female specimens which have a lot of black on the head like the
males, but specimens occur too without any black markings, or even
with a reddish crown, which resembles the colour of the back."
(Schwarz, 1931, pp. 410, 412-413.)
"The range of this form is only imperfectly known. It clearly
inhabits the south-eastern coast from Ft. Dauphin in the south to
at least Masindrano in the north. How far it goes into the interior
is not known, but it would appear that it is restricted to the coastal
forests, as L. /. rufus is already found in eastern Betsileo. It also
remains to be ascertained where the ranges of collaris and julvus
meet." Specimens are recorded from: Farafangana, about 23 S. ;
Loholoka, about 21 60' S.; and Fort Dauphin. (Schwarz, 1931,
p. 412.)
Ten specimens of "Lemur nigerrimus" were collected by the
Mission Zoologique Franco-Anglo-Americaine of 1929-31 (Delacour,
1932, p. 220).
"At Vondrozo, in June and July, this lemur was common in the
rain forest, usually traveling in the trees in parties of from four
to six. . . .
"The animals were not shy and could be closely approached ....
"Several stomachs examined contained woody fruits of forest
trees and one, green vegetable matter that was possibly leaves.
"At Manomba in October many females were carrying young."
(Rand. 1935, p. 97.)
138 EXTINCT AND VANISHING MAMMALS
The same author describes the native method of capturing these
lemurs by means of snares arranged on "a line of poles set up on
forked sticks across a long, narrow clearing, forming a bridge from
the trees on one side of the clearing to those on the other."
[In view of the still existing uncertainty concerning the taxonomic
status of Prosimia flavijrons J. E. Gray, the exact type locality of
which is unknown, it seems hardly worth while to give a separate
account of it here. Schwarz (1931, p. 412) , after provisionally con-
sidering it a synonym of L. m. collaris, proposes later (1936, p. 24)
to recognize it as a distinct subspecies of L. macaco and assigns
to it a range at Maromandia, northwestern Madagascar. On the
other hand, G. M. Allen (1939, p. 127) regards it as "probably a
synonym of Lemur macaco collaris" whose range lies at approxi-
mately the opposite end of Madagascar from Maromandia.]
Red-fronted Lemur
LEMUR MACACO RUFUS Audebert
Lemur Rujus Audebert, Hist. Nat. Singes et Makis; Makis, p. 12, 1800. (Type
locality not stated = Madagascar.)
SYNONYM: Lemur rufifrons Bennett (1833).
FIGS.: Audebert, 1800, Makis, pi. 2; Milne Edwards and Grandidier, 1890,
pis. 138, 139, 145.
This is apparently one of the more common of the Madagascar
lemurs.
Female (type of rujus) : similar in size to L. mongoz, but differs
in shorter ears, shorter hairs on tail, and rufous pelage; snout, and a
line from forehead to crown, black; crown, temples, cheeks, and
throat dirty white; body yellowish rufous; tail brown at the tip
(Audebert, 1800, Makis, p. 12) . Male (type of "rufifrons") : back
grizzly; tail darker; under parts, haunches, and limbs mixed with
rufous; forehead and cheek-tufts rufous; a nearly complete circle
of white about the eye; nose, and line through middle of forehead,
black (Bennett, 1833, p. 106) . The male differs from males of other
subspecies of L. julvus in having a rusty-red forehead (Schwarz,
1931, p. 410). This form is very variable in coloration.
Schwarz (1931, pp. 413-414) describes its occurrence as follows:
The range of L. f. rujus includes the greater part of Madagascar, all the
central plateau, and the west coast. ... It would appear that, except the
north and a fairly narrow strip on the east coast, the whole mainland of
Madagascar is inhabited by this race.
As a matter of fact the majority of the individuals found north of the
Betsiboka River are black-headed and much like the black-headed phase of
L. /. albijrons; but there can be no doubt that a mixed population is found
in a considerable part of the north. At present I am not prepared to say
ORDER PRIMATES: PRIMATES 139
definitely whether this is due to mutation or, what is more probable, to
secondary invasion of the northern area by L. /. rujus down the right bank
of the Betsiboka River. Red-fronted skins have been recorded from north
of the River Betsiboka from Betsako . . . , Narendry Bay . . . , Anoront-
sanga . . . ; also . . . from Ambatondrazaka, south of Lake Alaotra. Black-
headed skins have been recorded from the same general region, but not south
of the River Betsiboka. In various cases black- and red-fronted skins are
known from the same localities.
Additional localities from which Schwarz records specimens are:
Ankona Forest, E. Betsileo; Fianarantsoa ; Tulear, SW. coast; and
Morondava, W. coast.
Lorenz-Liburnau (1898, p. 448) records 16 specimens from Kan-
dam and 3 from nearby Antema in the Bay of Bombetoka region.
In the forests in the vicinity of Betsina, west of the Mahavavy
River, northwestern Madagascar, this lemur is not rare, and four
specimens were secured (Kaudern, 1915, p. 45) .
G. M. Allen (1918, p. 515) records "a fine series of six males and
ten females, all from localities on the upper Siribihina River [inland
from Morondava] and some thirty miles south of Berevo."
"This lemur was common about Tabiky [inland from Cape St.
Vincent] in the low dense brush, the wooded plains, and the gallery
forest." On one occasion a party of six was seen. (Rand, 1935,
p. 98.)
Mayotte Lemur
LEMUR MACACO MAYOTTENSIS Schlegel
Lemur mayottensis Schlegel, Nederl. Tijdschr. Dierk., vol. 3, p. 76, 1866.
("L'ile de Mayotte," Comoro Isles.)
FIG.: Schlegel and Pollen, 1868, pi. 2.
In former years the Mayotte Lemur was apparently common, but
with the increase of population and cultivation on the island its
numbers have probably declined.
Coloration of the head similar to that of L. /. collaris but much
more pronounced; in old males the snout is deep black, this colora-
tion extending between the eyes to the forehead and continuing as
a median stripe to the crown; this color pattern of the head less
pronounced in younger males and in females; upper parts grayish
brown, speckled with rufous and yellowish gray; rump with a
blackish spot; lower parts pale rufous (Schlegel, 1866, pp. 76-77).
Total length, 39 inches; tail, 21 inches (Schlegel, 1876, p. 308).
These animals live in bands of 6 to 20 individuals in the virgin
forests of Mayotte, where they are hunted with dogs. The flesh is
excellent and tastes like that of young rabbits. A favored food of
the lemurs is wild dates. A series of 10 specimens is recorded.
(Schlegel and Pollen, 1868, pp. 5-6.)
140 EXTINCT AND VANISHING MAMMALS
Mongoose Lemur
LEMUR MONGOZ MONGOZ Linnaeus
[Lemur] Mongoz Linnaeus, Syst. Nat., ed. 12, vol. 1, p. 44, 1766. (Based upon
the "Mongooz" of Edwards (1758, p. 12); type locality, "Madagascar.")
SYNONYMS : Lemur nigrijrons, L. albimanus, and L. anjuanensis of fi. Geoffroy-
Saint-Hilaire (1812) ; Prosimia micromongoz, P. bugi, P. brissonii, and
P. ocularis of Lesson (1840) ; Lemur cuvieri Fitzinger (1870) ; Propithecus
brissonianus J. E. Gray (1870).
FIGS.: Edwards, 1758, pt. 1, pi. 216; Schreber, Saugthiere, vol. 1, pis. 39 B, 42
("Lemur Simia-Sciurus") , 1774; Audebert, 1800, Makis, pi. l re ; Geoffrey
and Cuvier, Hist. Nat. Mamm., livr. 2, pi. 11, 1819, livr. 30, pi. 176, 1821;
Proc. Zool. Soc. London 1871, pi. 16; Milne Edwards and Grandidier,
1890, pis. 156, 157, 162, 163, 164, 165 (figs. 1, 2); Elliot, 1913, vol. 1, pi. 4,
lower fig. ("Lemur nigrijrons").
In 1915 this lemur was reported as found now and then in the
forests of northwestern Madagascar, but as apparently nowhere
common (Kaudern, 1915, p. 43). Concerning its present numerical
status on certain islands of the Comoro group we have no in-
formation.
Size less than that of a small cat; area about eyes and tip of
nose black ; area beneath eyes white ; upper parts dark brownish ash
color; under parts white; tail long (Edwards, 1758, pt. 1, p. 12).
A white- and a red-cheeked phase, generally sex-linked ; anal region
almost naked. "In the red-cheeked phase there is not the black
crown-patch found in the male of L. m. coronatus, whereas in the
white-cheeked phase the colour of the cheeks is brighter than in the
female of L. m. coronatus, and a large black or blackish crown-
patch developed which is absent in that race. The tail is grey or
blackish in both sexes in mongoz; in coronatus it is reddish in the
female. As far as my experience goes the males invariably have
red and the females white cheeks in all specimens from Anjouan
and Moheli, Comoro Island [s]." (Schwarz, 1931, pp. 414-415.)
"This race is found on the Comoro Islands, Anjouan and Moheli,
but not on Mayotte .... On the mainland of Madagascar this
form is found on the south bank of the Betsiboka River, which it
ascends up to its head-waters." Specimens are recorded from:
Anjouan and Moheli Islands; Antema, Bay of Bombetoka, S. bank;
and Ambatondrazaka, south of Lake Alaotra. (Schwarz, 1931,
p. 416.)
Lorenz-Liburnau (1898, pp. 450-451) records 14 specimens (as
L. albimanus} from Kandani, and 5 from Antema, on the south side
of the Bay of Bombetoka.
G. M. Allen (1918, p. 515) records a specimen from Didy, soufh
of Lake Alaotra.
Petit (1931, p. 560) records a specimen from Ambongo, north-
western Madagascar.
ORDER PRIMATES: PRIMATES 141
Delacour (1932, p. 219) records 18 specimens of "Lemur nigri-
frons" collected by the Mission Zoologique Franco-Anglo-Ameri-
caine of 1929-31.
Crowned Lemur
LEMUR MONGOZ COBONATUS J. E. Gray
Lemur coronatus J. E. Gray, Ann. Mag. Nat. Hist., ser. 1, vol. 10, no. 65,
p. 257, 1842. ("Madagascar"; type locality restricted by Schwarz (1931,
p. 416) to "Bay of Mahajamba, N.W. coast (15 14' S.).")
SYNONYM: Lemur chrysampyx Schuermans (1848).
FIGS.: J. E. Gray, Zool. Voy. Sulphur, Mammalia, pi. 4, 1844; Schuermans,
Mem. Couronnes et Mem, Savants Strangers, Acad. Roy. Belgique, vol. 2,
pi. facing p. 6, 1848; Milne Edwards and Grandidier, 1890, pis. 158-161,
165 (figs. 3, 4), 166.
This subspecies of northern Madagascar apparently still remains
very common.
"Ashy above, limbs and beneath pale yellowish; face white;
orbits gray; cheeks and forehead bright rufous, with a large black
spot on the crown; tail thick, end blackish" (J. E. Gray, 1842,
p. 257). Males with red, females with whitish cheeks; anal region
thickly haired; tail reddish in the female (Schwarz, 1931, pp. 414-
415).
"This race of L. mongoz is found in northern Madagascar, both
in the east and west, north of the bays of Bombetoka and Antongil
respectively." Specimens are recorded from: Vohemar, NE. coast;
Ampasimbato, Central N. Madagascar; Amber Mountains; and Bay
of Mahajamba. (Schwarz, 1931, p. 416.)
Lorenz-Liburnau (1898, p. 449) records one specimen from Betsako
and four from near-by Ambundube, north of the Bay of Bombetoka,
under the name of L. mongoz nigrijrons. According, however, to
Schwarz (1931, p. 416), the animals of this area are coronatus.
Delacour (1932, p. 219) reports 39 specimens collected by the
Mission Zoologique Franco-Anglo-Americaine of 1929-31. Judged on
this basis, it is one of the commonest lemurs of Madagascar.
"Very common in the dry wooded areas of the northern savanna,
sometimes in rather low dense brush ; found also in dry forest on the
slopes of Mt. d'Ambre, up to about 800 meters. It was absent,
however, from the humid forest on the summit. . . .
"The animals were very tame and the natives sometimes killed
them with sticks. . . .
"Near Vohemar, DuMont and I saw a party of seven ....
"At Tarakibany ... I saw a party of five." (Rand, 1935, p. 98.)
142 EXTINCT AND VANISHING MAMMALS
Red-bellied Lemur
LEMUR RUBRIVENTER I. Geoffroy-Saint-Hilaire
Lemur rubriventer I. Geoffroy-Saint-Hilaire, Compt. Rend. Acad. Sci. [Paris],
vol. 31, p. 876 (1850), 1851. ("Madagascar.")
SYNONYMS: Lemur flaviventer I. Geoffroy-Saint-Hilaire (1851); L. rujiventer
J. E. Gray (1870); Prosimia rufipes J. E. Gray (1871).
FIGS.: Milne Edwards and Grandidier, 1890, pis. 167-170; J. E. Gray, Proc.
Zool. Soc. London 1872, pi. 69.
Even 30 years ago this species was considered rather rare on the
eastern coast of Madagascar (Kaudern, 1915, p. 47) .
It may be distinguished from all the other lemurs by the rufous-
chestnut of its under parts, limbs, and ruff; upper parts speckled
rufous-brown; tail blackish (I. Geoffroy-Saint-Hilaire, 1851, p. 876).
Elliot (1913, vol. 1, p. 152) adds the following details from the type
(a male) in Paris: line from forehead, top of nose, and lips maroon;
head above mixed dark brown and buff; body above chocolate-
brown; base of tail maroon, rest blackish; total length, 711 mm.;
tail, 407 mm. Schlegel's measurements (1876, p. 311) are larger:
total length, 38 inches; tail, 20 inches.
Schwarz (1931, p. 417) records specimens from the following
localities: Vohemar, NE. coast; Bay of Antongil; Betsimisaraka
country, west of Tamatave; Tamatave, NE. coast; Forest of Ankay,
NE. of Tananarive; Ambohimitombo and Ivohimanitra, N. Tanala
country; Vinanitelo, "SW." [=SE.] Betsileo; Manakara River, SE.
coast; Mojanga, Bay of Bombetoka; and Morondava, W. coast. "If
all these records are correct the range of L. rubriventer includes the
greater part of Madagascar, not only the eastern forest region, as
has hitherto been supposed."
Delacour (1932, p. 219) records 21 specimens collected by the
Mission Zoologique Franco-Anglo-Americaine of 1929-31.
"This diurnal lemur was found in small parties similar to the
groups of L. julvus. One party contained at least four adults and
five young. To the west of Andapa [in northeastern Madagascar]
. . . eight young . . . were taken with the adults." (Rand, 1935,
p. 98.)
Ruffed Lemur. Vari (Fr.)
LEMUR VARIEGATUS Kerr
L[emur] Macaco variegatus Kerr, Anim. Kingdom of Linnaeus, p. 86, 1792.
(Based upon the "Vari" of Smellie's Buffon (vol. 7, pi. 229, 1791?); type
locality, "the islands of Madagascar and Johanna, and the neighbouring
countries of Africa" = Madagascar.)
SYNONYMS: Lemur ruber 1C. Geoffroy-Saint-Hilaire (1812); Prosimia erythro-
mela Lesson (1840); Lemur varius I. Geoffroy (1851).
ORDER PRIMATES: PRIMATES 143
FIGS.: Buffon, Hist. Nat., vol. 13, pi. 27, 1765; Schreber, Saugthiere, vol. 1,
pi. 40 B, 1774; Audebert, 1800, Makis, pis. 5, 6; Gervais, Hist. Nat. Mam-
mif., pt. 1, pi. 10, 1854; Milne Edwards and Grandidier, 1890, pis. 123-129;
Forbes, 1894, vol. 1, pi. 7; Beddard, 1902, p. 542, fig. 259; Elliot, 1913,
vol. 1, pi. 5, lower fig. (facing p. 158) ; Kaudern, 1915, pi. 3, fig. 1.
This lemur is apparently still common in northeastern Mada-
gascar.
"Mostly white in the body; all the paws, the fore-head, the
tail, the insides of the thighs, a large blotch on each shoulder, and a
long narrow patch on the loin before the upper part of the thigh,
are black .... The muzzle is long and thick, the ears very short,
and fringed with long flowing hairs, which join the collar, or hairy
ruff, on the neck, cheeks, and throat." (Kerr, 1792, pp. 86-87.)
A color phase, described as "Lemur ruber," has a general rufous
color; head, hands, tail, and abdomen black; a half-collar of white
on top of the neck (E. Geoffroy-Saint-Hilaire, 1812, p. 159).
Total length 44 inches, of which the tail makes up nearly half
(Schlegel, 1876, p. 302).
Schwarz (1931, p. 418) records specimens from: Tombato River;
Sakana, opposite the He Ste. Marie; Alumanitra Forest; Bay of
Antongil; and Ambatondrazaka, south of Lake Alaotra. He adds:
"Black and white and red specimens have been collected in the
same place by J. Audebert in the coast region north of the Bay of
Antongil. Red specimens are recorded from Maroansetra, Bay of
Antongil; Malewo and Andranofotsy, N.E. coast, north of the Bay
of Antongil. . . .
"From the material at hand it would thus appear that Lemur
variegatus is restricted to the forests of N.E. Madagascar. Its
northern limit may be about 13 30' S., its southern range has been
ascertained as the region of Tamatave at about 18 S. . . . On the
plateau Ambatondrazaka remains the only place known."
Coquerel writes (1859, p. 462) that this animal is sacred to the
inhabitants of Tamatave; they say that it worships the sun and
prays to it every morning.
Pollen stated (1868, p. 21) that up to that time it had been
observed only in the forests of the region between Tintingue, Tama-
tave, and Antananarivo. It was found there in considerable bands,
living on fruits.
Kaudern (1915, p. 43) records specimens from west of Fenerive
and south of Tamatave.
G. M. Allen (1918, p. 516) records a specimen from 100 miles west
of Tamatave.
The Mission Zoologique Franco-Anglo-Americaine of 1929-31 col-
lected 8 specimens of variegatus and 9 of "ruber" (Delacour, 1932,
p. 219).
144 EXTINCT AND VANISHING MAMMALS
"Near Maroantsetra (two days northeast) individuals in the red
phase, the only phase seen there, were common. The variegated
lemur is diurnal and arboreal, usually seen in pairs." (Rand, 1935,
p. 99.)
Weasel Lemur
LEPILEMUR MUSTELINUS I. Geoffrey
L[epilemur] mustelinus I. Geoffrey, Cat. Method. Mamm. [Mus. Paris], pt. 1,
Primates, p. 76, 1851. ("Madagascar"; type locality restricted by Schwarz
(1931, p. 420) to "Tamatave.")
SYNONYMS: Mixocebus caniceps Peters (1875); Lepidolemur microdon Major
(1894).
Fios.: Peters, Monateb. Preuss. Akad. Wiss. Berlin 1874, pi. 1 (facing p. 694),
1875 (as Mixocebus caniceps) ; Milne Edwards, Grandidier and Filhol,
1897, pi. 255, pi. 259, fig. 1.
Only four specimens of this species were collected by the Mission
Zoologique Franco-Anglo-Americaine of 1929-31 (Delacour, 1932,
p. 220) . This fact, in connection with the meager information con-
cerning the animal, indicates that it is one of the less common of
the Madagascar lemurs.
General color rufous; throat white; forehead and cheeks gray;
under parts and inner side of limbs yellowish gray; last third of
tail brown; rest of tail and lower part of limbs yellowish gray.
Head and body, about 350 mm.; tail, 250 mm. (I. Geoffrey, 1851,
p. 76.)
"The range of L. mustelinus includes the moist east and north-
east of Madagascar, at least as far south as Betsileo and as far
north as Vohemar. It does not occur in the north-west, where L.
ruficaudatus is found." Specimens are recorded from: Vohemar;
Ankay Forest, NE. of Tananarive; Ampitambe and Antsiraka,
Betsimisaraka country; Ankona Forest; Upper Masiatra River, E.
Betsileo; and Vinanitelo, "SW." [ = SE.] Betsileo. (Schwarz, 1931,
p. 420.)
G. M. Allen (1918, p. 516) records a specimen from Didy, south
of Lake Alaotra.
All the localities of this species are on the eastern slope of the
island, from Vohemar on the north to Vinanitelo on the south ; these
localities are littoral or belong to the region of the Hauts-Plateaux.
The altitudes where it is found vary from less than 100 m. to more
than 1,000 m. It is probably divisible into subspecies not yet de-
termiried. All alleged records from the west coast really belong to
L. ruficaudatus. (Petit, 1933, p. 34.)
ORDER PRIMATES: PRIMATES 145
Red-tailed Lemur. Hattock (Madagascar)
LEPILEMUR RUFICAUDATUS Grandidier
LepilemMr ruficaudatus Grandidier, Rev. Mag. Zool., ser. 2, vol. 19, p. 256,
1867 - 1 ("La cote sud-ouest de Madagascar"; type locality restricted by
Elliot (1913, vol. 1, pp. 122) to "Morondava, Madagascar.")
SYNONYMS: Lepilemur dorsalis Gray (1870); L. pallidicauda Gray (1873);
Lepidolemur leucopus Major (1894); L. edwardsi Major (1894); L.
globiceps Major (1894) ; L. grandidieri Major (1894) ; L. mustelinus
rufescens Lorenz-Liburnau (1898).
FIGS.: Forbes, 1894, vol. 1, pi. 9 (as L. leucopus}] Milne Edwards, Grandidier
and Filhol, 1897, pi. 256 (as L. m. var. dorsalis), pi. 257, pi. 258 (as L. m.
var. leucopus), pi. 259 (as L. dorsalis); Lorenz-Liburnau, 1898, pi. 30 (as
L. m. rufescens); Kaudern, 1915, pi. 1, fig. 3, pi. 2, fig. 2 (as L. m.
rufescens) .
This is apparently a rather common as well as widespread species
in western and southern Madagascar. No less than 36 specimens
were collected by the Mission Zoologique Franco-Anglo-Americaine
of 1929-31 (Delacour, 1932, p. 220) .
General color ashy rufous; head blackish; hind limbs pale ashy;
tail reddish; throat fulvous; abdomen whitish. Total length, 560
mm.; tail, 250 mm. (Grandidier, 1867, p. 256.)
Schwarz (1931, pp. 420-421) records specimens from the following
localities: Loko-Be, Nosy Be Island, NW. Madagascar; Betsako,
north bank, Bay of Bombetoka ; Ambundube, near Betsako ; Antema
and Kandani, south bank, Bay of Bombetoka; Morondava, W. coast;
Ambolisatra, SW. coast; and Fort Dauphin, SE. coast. "This species
has also been recorded by Pollen and van Dam from various points
on the N.W. coast: Anorontsanga, Jangoa River, Ampasindava, all
at or near the Bay of Ampasi[n]dava; also from the Bay of Maha-
jamba. The range, therefore, covers the entire west coast, including
the north-west and south-east, but not the central plateau. The
northern and eastern limits remain to be ascertained." (P. 421.)
Schlegel and Pollen write (1868, p. 12) concerning this species
(under the name of L. mustelinus) that it is very stupid and more
slothful than Hapalemur griseus. The natives of the northwest say
that they sometimes kill it in daytime with sticks and eat its flesh.
It is nocturnal, and its food consists of buds and leaves of trees as
well as certain fruits.
Kaudern (1915, p. 74) records several specimens from Ste. Marie
de Marovoay on the Betsiboka River, northwestern Madagascar.
1 Petit (1933, pp. 36-37) recognizes three subspecies, including L. r. dorsalis
Gray and L. r. leucopus Major. However, he attempts to distinguish dorsalis
from ruficaudatus in part by "a more russet coloration" and by "a constant
and more distinct dorsal band," in contradiction to Gray's type description of
dorsalis (1870, p. 135): "Back grey, yellow-washed; dorsal stripe none." As for
leucopus, Schwarz states (1931, p. 420) that "the type-specimen is in every
respect typical ruficaudatus"
6
146 EXTINCT AND VANISHING MAMMALS
In recognizing dorsalis as a subspecies, Petit states (1933, p. 37)
that it inhabits the Mahafaly and Antandroy districts, and dis-
appears abruptly with the very distinct limit of the xerophytic
vegetation near Bevilana, west of Fort Dauphin. The same author
(1935, p. 474) remarks on its former presence in the Manampetsa
Reserve in the southwest, but it was not found there in 1932 or 1933.
Rand (1935, p. 99) records the "Hattock" from Tabiky and
Tsarakibany.
Family INDRIIDAE: Sifakas, Indri, Avahis
The Sifakas and their relatives constitute a family of 3 genera
and 13 forms. All are endemic to Madagascar. Some of the natives
have a certain degree of superstitious veneration for the elegantly
attired Sifakas and the Indri, and yet the animals are not altogether
free from persecution. In numerical status they vary from common
to rare. Their limited distribution and their uncertain future render
all of them suitable subjects for inclusion in the present report.
Diademed Sifaka. Propitheque a diademe (Fr.)
PROPITHBCUS BIADEMA DIADBMA Bennett
Propithecus Diadema Bennett, Proc. Zool. Soc. London 1832, p. 20, 1832.
("Madagascar"; type locality restricted by Milne Edwards and Grandidier
(1875, p. 300) to "la cote Est de Madagascar." 1 )
SYNONYMS: Macromerus typicus A. Smith (1833); Indris olbus Vinson (1862).
FIGS.: Gervais, Hist. Nat. Mammif., pt. 1, pi. 8, 1854; Milne Edwards and
Grandidier, 1875, pi. 1.
Over a hundred years ago this Sifaka was said to be rare, and
it is apparently still more so today. The Mission Zoologique Franco-
Anglo-Americaine of 1929-31 collected only three specimens the
smallest number reported for any of the Madagascar lemuroids
(Delacour, 1932, p. 220) .
Face nearly naked; hairs generally long, silky, waved, erect, and
glossy; ears rounded, concealed within the fur; a yellowish-white
band extending across the forehead and below the ears to the throat;
crown, nape, and hands black; shoulders, sides, and lower back
mixed black and white; limbs, rump, and tail pale fulvous; throat
like sides, rest of under parts white. Head and body, 21 inches;
tail, 17 inches. (Bennett, 1832, pp. 20-21.)
This subspecies is found only between the Bay of Antongil on the
north and the Masora River on the south [at about lat. 20 S.],
i Elliot (1913, vol. 1, pp. 169, 171) attempts the impossible in stating that
Sambava, northeastern Madagascar, is the type locality of this subspecies as
well as of P. d. sericeus. For the latter he also attempts to switch Sambava
to the northwest coast.
ORDER PRIMATES: PRIMATES 147
in the narrow bands of forests on the eastern slope of the moun-
tains (Milne Edwards and Grandidier, 1875, p. 300) .
"It is stated to be rare" (Bennett, 1832, p. 22).
G. M. Allen (1918, p. 515) records a specimen from Didy, south
of Lake Alaotra.
Schwarz (1931, p. 422) mentions specimens from: Tamatave,
NE. coast ; Andragoloaka, SE. of Tananarive ; and Mananare, Bay of
Antongil.
Silky Sifaka. Propitheque soyeux (Fr.)
PROPITHECUS DIADEMA CANDIDUS Grandidier
Propithecus candidus Grandidier, Compt. Rend. Acad. Sci. [Paris], vol. 72,
p. 232, 1871. ("Les forets au nord de la baie d'Antongil, sur la cote est
de Madagascar"; type locality restricted by Schwarz (1931, p. 421) to
"Sahambavany, N.E. coast" of Madagascar.)
SYNONYM: Pr&pithecus sericeus Milne-Edwards and Grandidier (1872).
FIG. : Milne Edwards and Grandidier, 18756, pi. 2.
This Sifaka has a limited range in the northeast of Madagascar,
and its numbers are apparently few.
It is distinguished from P. verreauxi by its entirely white color,
without black crown or ashy spot on the back (Grandidier, 1871,
p. 232). Muzzle bare, black, with spots of flesh color; pelage gener-
ally entirely white, with silky reflections and washed lightly with
yellow; hairs at base of tail and beneath claws rufous; various
intergradations between this subspecies and P. d. diadema observed
(Milne Edwards and Grandidier, 1875a, p. 301). Head and body,
620 mm.; tail, 400 mm. (Milne-Edwards and Grandidier, 1872,
p. 274).
Schwarz (1931, pp. 421-422) records specimens from Sahamba-
vany, NE. coast, and from Antsompirina and Ansandrizina, NE.
coast (probably on the peninsula which forms the eastern border
of the Bay of Antongil). "The range of this race includes the
north-eastern coast ranges of the island from the region of Bemarivo
(14 16' 30" S.), which is the northernmost locality recorded, to the
Bay of Antongil ; I suppose the Tsingambala River, at the northern
end of the bay, will be found to separate the range of candidus from
that of diadema."
The Silky Sifakas inhabit the narrow bands of forests covering
the eastern slope of the mountains in the northeast, between the
Lokoy River (13 miles south of Sambava) and the Bemarivo River
(10 miles north of Sambava). They go ordinarily in smaller bands
than their congeners; scarcely more than three or four are found
together. (Milne Edwards and Grandidier, 1875a, p. 302.)
Elliot (1913, vol. 1, p. 171) is in error in placing the range in
northwestern, instead of northeastern, Madagascar.
148 EXTINCT AND VANISHING MAMMALS
The Mission Zoologique Franco-Anglo-Americaine of 1929-31 col-
lected six specimens (Delacour, 1932, p. 220) .
Milne-Edwards's Sifaka. Propitheque d'Edwards (Fr.)
PROPITHECUS DIADEMA EDWARDSI Grandidier
Propithecus Edwardsi Grandidier, Compt. Rend. Acad. Sci. [Paris], vol. 72,
p. 232, 1871. ("Les forets situees dans Pouest de Mananzary, a Mada-
gascar" ; type locality further delimited by Milne Edwards and Grandidier
(1875a, p. 303) as "la foret de Manampahy," in southeastern Madagascar.)
SYNONYM: Propithecus bicolor Gray (1872).
FIG.: Milne Edwards and Grandidier, 18756, pi. 3.
No information is at hand concerning the numerical status of
this subspecies.
It is entirely black, save for an area of rufous-white on each
side of the loins; face naked and black; ears well developed and
covered with long hairs. Head and body, 640 mm.; tail, 460 mm.
(Grandidier, 1871, p. 232.) Milne Edwards and Grandidier (1875a,
p. 303) give additional details: black areas lightly washed with
rufous; a narrow band of reddish black separating the light lumbar
areas; a light reddish spot at the base of the tail; considerable
variation in coloration evident, some individuals exhibiting an ap-
proach to P. d. diadema, and others to P. d. holomelas.
This Sifaka inhabits the narrow bands of forests that partly
cover the * eastern slope of the mountains between the Rivers
Masora (about 20 S.) and Matitanana (about 22 S.) (Milne
Edwards and Grandidier, 1875a, pp. 304-305) .
Schwarz (1931, p. 422) is in error in placing the type locality east
of Masindrano, which would be somewhere in the Indian Ocean.
He records specimens from: Ampitambe, Betsimisaraka country;
Ivohimanitra, Tanala; Ambohimotombo, N. Tanala; and Vinani-
telo, "S.W." [=S.E.] Betsileo. "The specimens from Vinanitelo
would indicate a considerable extension of the range on the central
plateau."
Black Sifaka
PROPITHECUS DIADEMA HOLOMELAS Giinther
Propithecus holomelas Giinther, Ann. Mag. Nat. Hist., ser. 4, vol. 16, p. 125,
1875. ("Fienerentova" Fianarantsoa, central Betsileo, Madagascar.)
Very little information about the Black Sifaka has ever been
assembled.
Nearly as large as P. d. edwardsi. "Throat and all the lower
parts covered with dense fine woolly hair. Male with a small patch
of ferruginous hairs ... in the middle of the chest . . . ; in the
ORDER PRIMATES: PRIMATES 149
female this patch is replaced by two smaller ones ... of a whitish
colour. All the upper parts deep black, except the back of the root
of the tail, which is brownish. Abdomen greyish brown. A few
whitish hairs at the extremity of the tail." Head and body, 23 inches;
tail, 15-16 inches. (Glinther, 1875a, p. 125.)
Schwarz (1931, p. 423) records specimens from: Fianarantsoa ;
S. Betsileo; Nandihizana, N. Betsileo; and "Ambavombe," south
[ = west?] of Fort Dauphin, S. coast. "The range of this race
appears to be the inland mountain range in the south-east of
Madagascar, whereas P. d. edwardsi inhabits the coast range. I have
little doubt as regards the distinctness of the two races."
Verreaux's Sifaka. Propitheque de Verreaux (Fr.)
PROPITHECUS VERREAUXI VERREAUXI Grandidier
Propithecus Verreauxi Grandidier, Rev. Mag. Zool., ser. 2, vol. 19, p. 84, 1867.
("Les contrees arides et sablonneuses ou habitent les Antandroui's, les
Mahfales, et les Antitenes"; type locality restricted by Schwarz (1931,
p. 424) to "Tsifanihy, Prov. Antandroy, north of Cape Ste. Marie, S.
Madagascar." "Mananzari," erroneously stated by Elliot (1913, vol. 1,
p. 172) to be the type locality, is outside the entire range of the species.)
FIGS.: Grandidier, Album de Tile de la Reunion, vol. 4, pis. 1, 2, 1867;
Milne Edwards and Grandidier, 18756, pis. 4, 8.
This Sifaka has been reported as common in Madagascar from
Flacourt's time (1661) to the present.
It is similar to P. d. diadema but with a smaller body, longer
tail, and more whitish pelage; crown and nape rufous-brown;
shoulders and sides yellowish white; a gray dorsal patch often
present; limbs and hands white; face black, almost naked, with
some white hairs; under parts and inner side of limbs white; tail
white except at base; throat with a small longitudinal rufous spot.
Head and body, 490-500 mm.; tail, 450-600 mm. (Grandidier,
1867a, p. 84.)
Schwarz (1931, p. 424) records specimens from: Tulear, SW.
coast; Morondava, W. coast; Bemamanga near Morondava; Anta-
nosy country; and Ankazoabo, central SW. Madagascar. "Accord-
ing to A. Milne-Edwards and Grandidier the range of P. v. ver-
reauxi includes the whole south-western part of Madagascar from
the Tsidsobon River on the west coast to the region of Ft. Dauphin
in the south-east. Nothing is known of the exact limits of the range,
neither in the east, where it should meet P. d. holomelas, nor in
the west, where no record exists for the region between the Manan-
bolo and Tsidsobon Rivers, where either this race or P. v. deckeni
should occur."
Flacourt (1661, p. 153) reported many of these Sifakas in the
150 EXTINCT AND VANISHING MAMMALS
region west of Fort Dauphin. More than two centuries then elapsed
before they were rediscovered and named by Grandidier.
They inhabit the arid coasts in the south and the southwest, from
Andrahoumbe to the Tsidsibon River. They are always found in
bands of 10 to 12 individuals. They are diurnal and feed upon young
shoots of trees and upon fruits. (Grandidier, 1867a, p. 84, and
1867c, p. 313.)
In 1866 Grandidier encountered this Sifaka some leagues north
of the village of the Antandroy king, Tsifanihy, in the vicinity of
Cape Ste. Marie. Here he found it regarded with a certain venera-
tion by the natives; they objected to his skinning and dissecting a
specimen in their village, and they buried the remains ceremoniously
at some distance from the village. Later he secured a large number
at various points on the southwestern coast, especially along the
Morondava River, and at one time he had 15 living individuals. The
natives capture them by means of snares in the clearings. The ani-
mals never lived long in captivity. (Milne Edwards and Grandidier,
1875a, pp. 308-311.)
"This species was common on the Upper Siribihina River [inland
from Morondava], and at a locality twenty-five kilometers east of
Tulear" (G. M. Allen, 1918, p. 515).
Petit (1931, p. 559) records a specimen from the vicinity of An-
droka, in the Mahafaly country.
The Mission Zoologique Franco-Anglo-Americaine of 1929-31
collected 18 specimens (Delacour, 1932, p. 220) .
"The brown-capped Propithecus was very common in the wooded
areas of the southwest. A diurnal, arboreal creature, it was usually
seen in parties of five to eight. . . .
"They were usually not at all wild and could be watched at close
range. . . . Near Tulear ... a group in the trees by the roadside"
was noticed.
"Green leaves of trees are probably their staple food. . . . The
animals were often seen to feed on leaves and the young one I kept
for a time ate them eagerly." (Rand, 1935, pp. 100-101.)
This Sifaka has become rare in the Manampetsa Reserve in the
southwest (Petit, 1935, p. 474) .
CoquerePs Sifaka. Propitheque de Coquerel (Fr.)
PROPITHECUS VERREAUXI COQUERELI A. Milne-Edwards
Propithecus Coquereli A. Milne-Edwards, in Grandidier, Rev. Mag. Zool.,
ser. 2, vol. 19, p. 314, 1867. ("Nord-est" = probably northwestern Mada-
gascar.)
SYNONYM: Propithecus damonis J. E. Gray (1870).
FIGS.: Milne Edwards and Grandidier, 18756, pi. 6; Forbes, 1894, vol. 1,
pi. 11; Kaudern, 1915, pi. 1, fig. 1.
ORDER PRIMATES: PRIMATES 151
Up to about 30 years ago, Coquerel's Sifaka was very common
in parts of northwestern Madagascar (Kaudern, 1915, p. 4). On
the other hand, not a single specimen seems to have been taken by
the Mission Zoologique Franco-Anglo-Americaine of 1929-31 (Dela-
cour, 1932, p. 220).
General color white; a band of reddish brown extending across
the chest and onto the arms ; a spot of the same color on the upper
part of the legs; crown, hands, and inner side of the limbs pure
white; face covered with very short white hairs; tail slender; fur
woolly. Head and body of the type (a young individual) , 250 mm. ;
tail, 220 mm. (A. Milne-Edwards, in Grandidier, 1867c, p. 314.)
Adults have the head, cheeks, nape, and back yellowish white ; loins
dark reddish gray; sides and pelvic region dirty white; outer
side of arms and anterior side of thigh dark chestnut-rufous; tail
reddish gray (Milne Edwards and Grandidier, 1875a, p. 315) . Total
length of adults, 3 feet 4 inches (Schlegel, 1876, p. 293) .
This Sifaka is found only on the northwest coast of Madagascar,
from the south side of Narinda Bay to the north side of the Bay of
Bombetoka, between the Rivers Loza and Betsiboka. Numerous
specimens from this region have been examined. (Milne Edwards
and Grandidier, 1876, pp. 315-316.)
Lorenz-Liburnau (1898, p. 454) records five specimens from Bet-
sako and two from near-by Ambundube, north of the Bay of Bom-
betoka.
The animal is very common in the forests on the Ankarafantsika
Plateau, where it wanders about in bands of 3 to 10 or 12 individuals.
It was observed in several places between the Betsiboka and Maha-
jamba Rivers. At Ste. Marie de Marovoay, on the Betsiboka, several
hundred were seen in the wild, and about 60 specimens were shot.
(Kaudern, 1915, p. 4, map, p. 5.)
Additional specimens from Island Nosy Komba and from Am-
batondrazaka, south of Lake Alaotra, are recorded by Schwarz
(1931, p. 423). "The specimen from Ambatondrazaka . . . shows
an eastern extension of the range as far as Lake Alaotra."
Crowned Sifaka. Propitheque couronne (Fr.)
PROPITHECUS VERREAUXI CORONATUS A. Milne Edwards
P[ropithecus] coronatus "Pollen" A. Milne Edwards, Rev. Scientifique, ser. 2,
year 1, no. 10, p. 224, 1871. (Type locality not stated; restricted by Elliot
(1913, vol. 1, p. 174) to "Province of Boeny on the Bay of Bombetok,
Madagascar.")
SYNONYM: Propithecus damanus Schlegel (1876).
FIG.: Milne Edwards and Grandidier, 18756, pi. 7.
About 30 years ago the Crowned Sifaka was reported as common
(Kaudern, 1915, p. 6), but apparently none were taken by the
152 EXTINCT AND VANISHING MAMMALS
Mission Zoologique Franco-Anglo-Americaine of 1929-31 (Dela-
cour, 1932, p. 220) .
General color white; chest washed with brown; crown black (A.
Milne Edwards, 1871, p. 224). Similar in size to P. v. verreauxi;
forehead, head, and cheeks varying from dark blackish brown to
reddish gray; nape and remaining upper parts white, more or less
washed with rufous on the limbs and at the base of the tail; a gray
or brown spot on the nape ; tail and hands white ; under parts vary-
ing from light rufous to very dark rufous-brown (Milne Edwards
and Grandidier, 1876, pp. 318-319). Total length, 3 feet 5 inches;
tail, 21 inches (Schlegel, 1876, p. 294) .
This Sifaka inhabits the Boeny country, which is comprised be-
tween the sea on the north, the Betsiboka River on the east, and
the Manzaray [Mahavavy] River on the west. In the south some
were killed by Crossley not far from the great forest of Manerinerina,
where he secured a large number of Decken's Sifaka. (Milne Ed-
wards and Grandidier, 1876, p. 319.)
Lorenz-Liburnau (1898, p. 453) records 4 specimens from Antema
and 21 from Kandani.
A small band was seen in 1906 on the west side of the Betsiboka
River opposite Marovoay, and two specimens were collected in 1912
near the coast between the Bay of Bombetoka and the Mahavavy
River. It is not rare along this part of the coast, and it is very
common in the great forests on the Boeny Mountains. It does not
seem to occur east of the Betsiboka or west of the Mahavavy River.
(Kaudern, 1915, p. 6, map, p. 5.)
Decken's Sifaka. Propitheque de Decken (Fr.)
PROPITHECUS VERREAUXI DECKENII Peters
Propithecus Deckenii Peters, Monatsb. Preuss. Akad. Wiss. Berlin 1870, p. 421,
1871. ("Kanatzi [=Kanatsy], im 18 s. Br. an der Westkiiste von
Madagaskar" (Peters, 1869, p. 4).)
FIGS.: Peters, 1869, pi. 1 (as P. diadema) ; Milne Edwards and Grandidier,
18756, pi. 5.
This is still a common animal in western Madagascar.
The (immature?) type female is described by Peters (1871, p.
421) as having the hands and head yellowish white like the rest of
the body; lumbar region and sides washed with gray; face black,
with a whitish spot on the ridge of the snout; tail as long as, or
longer than, the head and body. Milne Edwards and Grandidier
add (1876, pp. 313-314) that adults have a little black diadem in
front of the white crown; upper chest bright rufous; rest of under
parts reddish white ; a tawny spot at the base of the white tail. Total
length 42 inches, of which the tail occupies half (Schlegel, 1876,
p. 295).
ORDER PRIMATES: PRIMATES 153
Crossley secured a number of specimens in the Forest of Manerine-
rina and in the plains north of Ankavandra [lat. 19 15' S.]. These
Sifakas inhabit the forests scattered here and there in the midst of
the great Jurassic plains lying between the Mananbolo and Manza-
ray [ = Mahavavy] Rivers. The Antimailaka natives consider them
sacred animals and never kill them. (Milne Edwards and Grandi-
dier, 1876, pp. 313-314.)
Decken's Sifaka is very common in the forests south of Lake
Kinkony (near the lower Mahavavy) , and five specimens were ob-
tained there. It is said not to occur in the forests between this lake
and Cape Tanjona. The Mahavavy River appears to form its north-
eastern limit. (Kaudern, 1915, pp. 6-7, map, p. 5.)
Beravina, 17 10' S., NW. coast, is one of the localities from
which Schwarz (1931, p. 424) records specimens.
No less than 30 specimens were collected by the Mission Zoologi-
que Franco-Anglo-Americaine of 1929-31 (Delacour, 1932, p. 220) .
"This Propithecus was common in the country from Namo-
roka to the Mahavavy Rivers, and a number apparently of the
same subspecies were seen between the Mahavavy and Betsiboka
Rivers. . . .
"This is a common, diurnal animal, found in parties of some-
times as many as nine individuals. It frequented the heavy gallery
forest, the lower, dryer forest, and at Soala I found a party in the
coast mangroves. . . . We found them rather tame here and easily
approached." (Rand, 1935, p. 99.)
Major's Sifaka
PROPITHECUS VERREAUXI MAJORI Rothschild
Propithecus majori Rothschild, Novit. Zool., vol. 1, p. 666, 1894. ("The
Antinosy country in south-west Madagascar" ; according to Schwarz (1931,
p. 424), this is the "country of the emigrated Antanosy, S. Central
Madagascar.")
FIG.: Rothschild, 1894, pi. 14.
Apparently no additional information has been secured concern-
ing this Sifaka since the original series was collected in 1889 and
described in 1894.
"Adult. Head and neck black. Face, snout, and ears naked,
and of a blackish colour, encircled by a broad band of long white
hairs, joining under the throat, slightly intermixed with darker
hairs. Rest of fur, including the tail, white on the upper surface,
back and upper rump dark brown. The large white patch on and
between the shoulders much grizzled with brown hairs. Upper sur-
face of hind limbs to just below the knees blackish brown. Inside
154 EXTINCT AND VANISHING MAMMALS
of hind limbs down to the heel also brown, joining the colour of the
upper surface, thus forming a continuous dark stripe along the
legs. Inner and upper surface of arms, thumb, and two following
fingers deep blackish brown; throat, chest, and greater part of
abdomen deep brown. Size perceptibly larger than that of Propi-
thecus verreauxi, with the tail longer.
"A number of specimens, all perfectly alike in colour, were sent
to me by Mr. Last from the Antinosy country in south-west Mada-
gascar. The collector also found Propithecus verreauxi Grandid.
in the same country, some with the back much darker than others,
but no specimens in any way intermediate between it and P. majori."
(Rothschild, 1894, p. 666.)
"There are four more skins exactly like the type in the Tring
Museum.
"The original label only says 'Antinosy' (=Antanosy) country.
It would appear that this means the mountain range round and east
of Manansoa (23 3' S., 44 50' E.), where J. T. Last was collecting
in 1889. The specimens of true verreauxi collected by Last and
also labelled 'Antinosy country' are probably from the plains farther
west, and collected on the way to or from Tulear, on the S.W. coast.
Only the original series of this race is known." (Schwarz, 1931,
p. 424.)
Perrier's Sif aka
PROPITHECUS PERRIERI Lavauden
Propithecus Perrieri Lavauden, Comptes Rendus Acad. Sci. [Paris], vol. 193,
no. 1, p. 77, 1931. ("Foret d'Analamera, situee au sud-est de Diego-
Suarez, dans le nord de Madagascar.")
This Sifaka is known only from a few specimens collected in the
Forest of Analamera, which covers an area of 5,000 or 6,000 hectares.
It bears some resemblance to Propithecus verreauxi coquereli of
western Madagascar but differs especially in its color. The entire
pelage is velvety black; ears small and glabrous; eyes brown. Head
and body, 500 mm.; tail, 450 mm.
It may seem surprising that this species has remained unknown
until so recently. The explanation lies in the fact that both the
animal itself and the forest in which it lives are strictly taboo to the
local Antakara natives. They give it the name of "Radjako"; this
was the name of a legendary hero among their ancestors. Few
Europeans have penetrated the Forest of Analamera. (Lavauden,
1931, pp. 78-79.)
G. M. Allen (1939t>, p. 133) regards the type specimen as "prob-
ably a melanistic individual of P. v. coquereli."
ORDER PRIMATES: PRIMATES 155
Inclri; Indris; Endrina. Indri (Fr.)
INDRI INDRI (J. F. Gmelin)
[Lemur} Indri J. F. Gmelin, Syst. Nat., vol. 1, pt. 1, p. 42, 1788. (Based upon
the "Indri" of Sonnerat (Voy. Indes or. et Chine, vol. 2, p. 142, 1782) ;
type locality, "Madagascar.")
SYNONYMS: Indris brevicaudatus E. Geoffroy (1796); Indri niger Lacepede
(1800) ; Indris ater I. Geoffroy (1825) ; Lichanotus mitratus Peters (1871) ;
Indris variegatus Gray (1872).
FIGS.: Sonnerat, 1782, vol. 2, pi. 88; Audebert, Hist. Nat. Makis, pi. 1, 1800;
Milne Edwards and Grandidier, 1875b, pis. 11, 12; Royal Nat. Hist., vol. 1,
p. 204, fig., 1893-94; Forbes, 1894, pi. 12.
The distribution of the Indri appears to be subject to considerable
local variation, but here and there in eastern Madagascar it is still
common.
This is the largest of the lemurs; it is almost entirely black; fur
silky and dense; snout, posterior under parts, back of thighs, and
lower arms grayish; rump white, with woolly hair; tail perceptible
only to the touch (Sonnerat, 1782, vol. 2, p. 142; J. F. Gmelin, 1788,
vol. 1, p. 42). It exhibits "a great variety of color pattern as well as
diversity of hues" (Elliot, 1913, vol. 1, p. 177). Head and body,
650 mm.; tail, 60-70 mm. (Milne Edwards and Grandidier, 1875a ;
p. 337).
Elliot (1913, vol. 1, p. 176) gives the range as "eastern coast of
Madagascar, in forests on the eastern side of the high mountains
between the Bay of Antongil on the north and the River Masara
on the south."
According to Sonnerat (1782, vol. 2, p. 142), the natives of the
south capture the animals when young, rear them, and train them
like dogs for hunting.
Pollen writes (1868, pp. 20-21) that up to that time the Indri
was known only from the interior of eastern Madagascar. His
friend Dr. Vinson reported that while passing through the great
forest of Alanamasoatrao he was deafened, during two days,' by the
incessant clamor of apparently numerous but invisible bands of
these animals. The natives have a superstitious veneration for the
Indri, and it plays quite a part in their folklore. It feeds upon fruits
and also preys upon small birds.
Milne Edwards and Grandidier state (1875a, pp. 340-341) that
the species lives only on the eastern slope of the great massif be-
tween the Bay of Antongil and the River Masora. It is essentially
diurnal and lives in bands, usually of no more than 4 or 5 indi-
viduals. They refute Sonnerat 's tale of its being trained by the
natives for hunting.
"This Lemuroid is probably the best known to travellers in
Madagascar, at least by ear, as no one can travel along the most
156 EXTINCT AND VANISHING MAMMALS
frequented route in the island, that from Tamatave to Antanana-
rivo, without often hearing the cries of these animals as he passes
through the great forest. They are not often seen." (Forbes, 1894,
vol. 1, p. 109.)
Schwarz (1931, p. 425) records specimens from the following
localities: Vohemar, NE. coast; Lalo River and Antsompirina, east
of the Bay of Antongil ; Sakana, opposite the He Ste. Marie ; Tama-
tave; Antsihanaka Forest, Lake Alaotra.
Specimens to the number of 16 were collected by the Mission
Zoologique Franco-Anglo-Americaine of 1929-31 (Delacour, 1932,
p. 219).
Rand writes as follows (1935, pp. 101-102) :
We encountered the indri only in the heavy forest of the northeast, from
sea level to 1800 meters, and found some surprising discrepancies in their range.
About the Bay of Antongil they were common forty kilometers northwest
of Maroantsetra, but at Maroantsetra, two days northeast, altitude 1000
meters, where the forest was equally heavy and continuous with that near
Maroantsetra, none were found. Again, east and north of Andapa there is
magnificent humid forest but none were heard in it, although west of Andapa,
perhaps eight kilometers away in similar forest continuous with the former,
they were common. . . .
The flesh of these creatures was well flavored but usually so tough and
hard even when the animals were fat that it was rather unsatisfactory food.
Many writers have said that this species is sacred to the Malagash. This
certainly is not true for the Malagash as a whole, for the people of the
south who had migrated to this part of the island had no objection to
skinning or eating these creatures, and even the native Betsimisaraka and
Tsimihity were quite ready to assist us in locating and shooting them, though
the Tsimihity at Andapa, one day west, would not eat the flesh.
Eastern Woolly Avahi. Avahis laineux oriental (Fr.)
AVAHI LANIGER LANIGER (J. F. Gmelhl)
[Lemur} laniger J. F. Gmelin, Syst. Nat., vol. 1, pt. 1, p. 44, 1788. ("Mada-
gascar." Lorenz-Liburnau (1898, p. 452) restricts this subspecies to the
east coast, and refers to Milne Edwards and Grandidier (1875a, 6), whose
pi. 9 represents a specimen from the Bay of Antongil. The vicinity of
this bay may be considered the restricted type locality.)
SYNONYMS: Lemur brunneus Link (1795); Lemur lanatus Schreber (1800?);
Indris longicaudatus fi. Geoffrey (1812) ; Semnocebus avahi Lesson
(1840); Avahis laniger orientalis Lorenz-Liburnau (1898).
FIGS.: Sonnerat, Voy. Indes or. et Chine, vol. 2, pi. 89, 1782; Schreber, Saug-
thiere, vol. 5, pi. 42 A, 1800(?); Gervais, Hist. Nat. Mammif., pt. 1, pi. 7,
1854; Proc. Zool. Soc. London 1866, pi. 15; Milne Edwards and Grandidier,
18756, pi. 9; Forbes, 1894, pi. 10; Elliot, 1913, vol. 1, col. pi. 7.
At the present time this appears to be one of the less common
of the Madagascar lemurs.
The hair is long and woolly, mouse gray at the base, reddish
brown in the middle, black at the tips; face broad, covered with
ORDER PRIMATES: PRIMATES 157
grayish-brown hairs; nose-pad alone nude; ears concealed, rufous;
a whitish band across the forehead, bordered anteriorly by a black
band ; back grayish brown ; a patch over the rump and region about
the base of the tail white, washed with rufous; under parts and
inner surface of limbs gray, washed with rufous; tail bright dark
red, deepest at its extremity (Forbes, 1894, p. 95). Body, 300 mm.;
tail, 390 mm. (Milne Edwards and Grandidier, 1875a, p. 327).
"A. I. laniger inhabits the whole forest region of north-eastern,
eastern, and south-central Madagascar." Specimens are recorded
from the following localities: Vohemar, NE. coast; Lakato Forest,
Ankay, NE. of Tananarive; Ambohitra, Kolaby Forest, N. Betsileo;
Vinanitelo, "S.W." [-S.E.], Betsileo; Fianarantsoa, central Bet-
sileo. (Schwarz, 1931, p. 426.)
According to Pollen (1868, p. 21), this species appears to be more
common than the Indri. It is recorded from the lie Sainte-Marie as
well as from the mainland of Madagascar.
The Avahi does not live in bands, but is always found singly or
in pairs. Its diet is exclusively vegetable. (Milne Edwards and
Grandidier, 1875a, p. 329; 1875b, map, pi. 122.)
"The first specimen . . . was brought to Europe by Sonnerat
. . . in 1781, and nearly half a century elapsed before a second one
was obtained" (Forbes, 1894, p. 96).
G. M. Allen (1918, p. 515) records a specimen from the Eastern
Forest.
The Mission Zoologique Franco-Anglo-Americaine of 1929-31
obtained only 9 specimens (Delacour, 1932, p. 219) .
Rand (1935, p. 102) records several individuals, including a party
of three, from the vicinity of Vondrozo in the southeast.
Western Woolly Avahi. Avahis laineux occidental (Fr.)
AVAHI LANIGER OCCIDENTALIS (Lorenz-Liburnau)
Avahis laniger occidentalis Lorenz-Liburnau, Abh. Senckenb. Naturf. Ges.,
vol. 21, p. 452, 1898. ("Ambundube," near Betsako, near Majunga, north-
western Madagascar.)
FIG.: Milne Edwards and Grandidier, 18756, pi. 10.
According to native report, this animal is not uncommon on the
Ankarafantsika Plateau and on the Bongolava of northwestern
Madagascar (Kaudern, 1915, p. 2).
Upper parts gray, with a yellowish-brown shade; woolly hair at
the base of the tail thin; tail reddish brown, toward the end more
blackish brown ; hands and feet yellowish brown ; face whitish ; un-
der parts cream-colored. Body, 330 mm.; tail, 195 mm. (Lorenz-
Liburnau, 1898, p. 452) .
Milne Edwards and Grandidier wrote (1875a, p. 329) that this
158 EXTINCT AND VANISHING MAMMALS
western form occurs between Mount d'Ambre and Anorontsangana,
near the Bay of Passandava. Since then the range has been extended
considerably to the southward.
"Collected by van Dam at Kakamba and Ampasidava, N.W.
coast, by Voeltzkow and [ = at] Ambundube near Betsako, and by
Kaudern from the Mahajamba River near Ste. Marie de Marovoay.
The range of this race includes the north-west coast as far south as
the Bay of Bombetoka; the northern and eastern limits are not
certain." (Schwarz, 1931, p. 427.)
Family DAUBENTONIIDAE: Aye-aye
The single representative of this family, the remarkable Aye-aye,
occurs in Madagascar, where it is decidedly rare.
Aye-aye
DAUBENTONIA MADAGASCARIENSIS (J. F. Gmelin)
[Sciurus] madagascariensis J. F. Gmelin, Syst. Nat., vol. 1, pt. 1, p. 152,
1788. (Based upon the "Aye-aye" of Sonnerat (Voy. Indes or. et
Chine, vol. 2, p. 137, 1782) ; type locality, "in occidental! parte insulae
Madagascar.")
SYNONYM: Lemur psilodactilus Schreber (Saugthiere, vol. 4, pi. 38 D, 1800?);
Cheiromys madagascariensis var. laniger G. Grandidier (1929).
FIGS: Sonnerat, Voy. Indes or. et Chine, vol. 2, pi. 86, 1782; Schreber,
Saugthiere, vol. 4, pi. 38 D, 1800?; Owen, 1863, pis. 14-19; Wolf, 1867,
pi. 3; Royal Nat. Hist., vol. 1, p. 241, fig, 1893-94; Forbes, 1894, pi. 1;
Beddard, 1902, p. 548, fig. 263; Lydekker, 1903, frontisp.; Elliot, 1907,
p. 552, fig. 80; EJliot, 1913, vol. 1, pi. 1; G. Grandidier, Bull. Acad.
Malgache, n. s., vol. 11, pi. facing p. 101, (1928) 1929.
The Aye-aye is perhaps the rarest as well as the most interesting
of all the surviving lemurs of Madagascar.
The head is short and round; patches of bristles above eyes and
nose and on cheeks and chin; eyes round, prominent; ears large,
rounded, naked, black; tail bushy, with hairs 3-4 inches long;
middle digit attenuated and wirelike. Fur on back, flanks, tail,
and limbs dark brown, nearly black; long hairs on top of head and
back of neck tipped with white; face, throat, under parts of body,
and inner side of limbs yellowish white; feet and digits black; tail
often with long white hairs throughout. Head and body, 18 inches;
tail, 18 inches. (J. F. Gmelin, 1788, p. 152, and Forbes, 1894,
pp. 14-16.)
"I have not seen any specimen with definite locality. The range
of this animal appears to include the whole forested portion of
Madagascar in the east, and apparently also in the north-west."
(Schwarz, 1931, p. 427.)
"I am told that the Aye-aye is an object of veneration at Mada-
ORDER PRIMATES: PRIMATES
159
gascar, and that if any native touches one, he is sure to die within
the year; hence the difficulty of obtaining a specimen. I overcame
this scruple by a reward of 10." (H. Sandwith, in Owen, 1863,
p. 38.)
This animal inhabits by preference the bamboo forests of the
interior. According to the natives, it is very rare; it lives solitarily
JE, .
FIG. 18. Aye-aye (Daubentonia madagascariensis) . From photo in Brehm.
or in pairs; and it is essentially nocturnal. It feeds on the pith of
bamboos and sugarcane and also on beetles and their larvae.
(Pollen, 1868, p. 22.)
"The Aye-aye lives in the dense parts of the great forest that
runs along the eastern border of the central plateau of the island,
but only in that part of it which separates the Sihanaka Province
from that of the Betsimisaraka, and which is about twenty-five
miles from the east coast, in latitude 17 22' S. or thereabouts. . . .
From what I have gathered from the natives, it seems to be pretty
common, its nocturnal habits and the superstitious awe with which
it is regarded . . . accounting for its apparent rarity ....
"Occasionally it is brought to Tamatave for sale, where it realizes
a good sum. Now and then it is accidentally caught in the traps
which the natives set for Lemurs." (Baron, 1883, pp. 639-640.)
160 EXTINCT AND VANISHING MAMMALS
"Many of the Betsimisaraka still believe that the Haihay is the
embodiment of their forefathers, and hence will not touch it, much
less do it an injury. It is said that when one is discovered dead in
the forest, these people make a tomb for it and bury it with all the
formality of a funeral." (G. A. Shaw, 1883, p. 45.)
"It was first discovered by Sonnerat during his travels in Mada-
gascar in 1780, and by him sent to Paris. The skin remained unique
in Europe for the best part of a century. ... It was for a long
period, and is still, very difficult to procure, or to induce the natives
to capture, specimens." (Forbes, 1894, pp. 16-17.)
Elliot (1913, vol. 1, p. 2) gives the range as "east coast from
Bay of Antongil to Mahanoro."
Kaudern (1915, p. 1) records four specimens from the forests west
of Fenerive and Tamatave on the east coast. He also mentions
(p. 2) some questionable reports of the species on the Ankara-
fantsika Plateau in the northwest.
G. M. Allen (1918, p. 516) records a specimen from Fenerive on
the Maningory River.
"The only aye-aye seen [by the Mission Zoologique Franco-
Anglo-Americaine of 1929-31] was in the northwest. Throughout
the rain forest of the east we found few who knew this creature,
but in the Sambirano it was well known to the natives by name,
though few had seen it. All of them said it fed on bamboo and was
very ferocious. . . . Our -single specimen was collected at Ampasa-
mena, a fishing village on the coast .... This individual ventured
into the village during the early part of the night and was walking
about amongst the houses when found by a native, who impaled it
on a fish spear. It was evidently not common or else not often seen
as the chief of the village, a gray-haired old man . . . , knew the
beast by name but had never seen one before." (Rand, 1935, p. 103.)
In view of the general tolerance and even awe exhibited by the
natives toward the Aye-aye, its rarity and possibly approaching
extinction must be attributable to more or less natural causes, as
yet undetermined.
Family COLOBIDAE: Leaf-eating Monkeys
The handsome Colobus Monkeys are externally distinguished
among African species by the reduction of the thumb, which is
either very small or altogether absent. A further point of structure
is in the sacculation of the stomach, a means probably for giving
greater capacity and a larger absorbing surface to the digestive
system, for the species are typically leaf-eaters and must in conse-
quence live upon a type of food requiring bulk and much digestion.
Two chief types occur, the black-and-white and the red groups.
ORDER PRIMATES: PRIMATES 161
These show much local variation, and many names have been ap-
plied. According to the latest reviser, Schwarz, however, these may
be regarded as representing but two distinct species, each with 19
or 20 races, or some 39 in all. They are typically monkeys of the
great rain forest, from French Guinea south to Angola, and across
the Congo Basin to the more isolated rain-forest and gallery forest
of Abyssinia (Ethiopia), Kenya, Tanganyika, and Zanzibar. Over
a large part of this range both species in one race or another occur
together, but in some regions only one of them is found, as in Zanzi-
bar, Kirk's Red Colobus, or in Abyssinia where the black-and-
white Guereza is alone represented. Both were first made known
from the West Coast, Sierra Leone.
G. M. A.
Black-and-white Colobus; Guereza
COLOBUS POLYKOMOS (Zimmermann) and races
Cebus polykomos Zimmermann, Geogr. Geschichte, vol. 2, p. 202, 1780.
(Sierra Leone.)
SYNONYMS and list of valid races: Schwarz (1929).
FIGS.: Elliot, 1913, vol. 3, pis. 3, 18, 19.
In these handsome black-and-white monkeys, the hair of the
flanks and hips tends to become elongate, the tail distinctly tufted,
with progressive increase in amount of white from all-black forms
(satanas) as in the Cameroons, to those with little and much white.
The most handsome of the races is perhaps kikuyuensis of Mount
Kenya or the race caudatus of Mount Kilimanjaro. For the char-
acters and synonymy of the various races, see Schwarz (1929).
On account of the long fine hair which forms the prominent
fringes along the sides, these monkeys are sometimes referred to as
"shawl monkeys." This quality seems also to have made them
desirable as fur so that a great many are killed. There is little in-
formation at hand as to the extent of this trade. Leplae (1925),
however, states that in the Belgian Congo the fur has a rather high
commercial value, and the species would be threatened with extinc-
tion if it were not protected by law. Such protection is given in the
British colonies but apparently not in the Congo to the extent that
it should be, although since 1929 it is given partial protection. In
Kenya Colony the race kikuyuensis occurs and on account of the
length of its white "shawl" is one of the handsomest of the races.
Its fur is, or not long ago was, much used by the natives in personal
decoration. Portions of the black-and-white fur are used as anklets
(particularly by the young men) or as caps. In the Gabun A. R.
Maclatchy (in litt., February 5, 1937) found them numerous in
bands in the mountainous region of Mimongo. They are of sedentary
habits and affect the high, abrupt mountains. "The vogue which
162 EXTINCT AND VANISHING MAMMALS
their magnificent skin enjoyed lately and even today has been the
cause of intensive hunting. A furrier of my acquaintance spoke of
having 30,000 skins in stock, collected from various parts of Africa
over several years. In view of the animal's restricted habitat, one
must admit that the protective decree was not unnecessary." These
monkeys are placed in Schedule B of the London Convention of 1933.
In general habits the Colobus Monkeys are in the main animals
of the dense saturate forests; they are not easy to find or shoot
and will often show considerable adroitness in hiding. Their food
consists largely of leaves, perhaps also lichens, among the hanging
festoons of which some of the races live, and probably small forest
fruits are also taken. Heller has recorded that in the Lado his party
came upon a troop of Colobus among thorn scrub, to which they had
come seeking the ripening bean pods, but on being approached they
made off over the ground to the nearest high forest. Such foraging
excursions must rather seldom be made in the case of the forms
which are more strictly high-forest dwellers. Apart from man, their
enemies are probably limited to leopards and the big crested eagles,
the food of which consists in part of monkeys.
While there seems to be little evidence that any of the races is
at present threatened with extinction, and since the demand for
their furs seems to have become less, they will no doubt be favored
by a limited permission to shoot specimens.
G. M. A.
Red Colobus
COLOBUS RADIUS (Kerr) and races
Simia (Cercopithecus) badius Kerr, Anim. Kingdom of Linnaeus, p. 74, 1792.
(Sierra Leone, based on the Bay Monkey of Pennant.)
SYNONYMS and list of valid races: See Schwarz, E., Zeitschr. f. Saugetierkunde,
vol. 3, pp. 92-97, June, 1928.
FIGS.: Elliot, 1913, vol. 3, pis. 5, 6, 14-16 (animal and skulls).
The Red Colobus Monkeys include no less than 20 recognized
races and differ in color from the black-and-white group, in having
the fur more or less black and red in varying pattern. Since the
fur is not as modified in long fringing patches along the sides, it is
not in special demand. Nevertheless one or two of the races are
rare or localized and may require special protection for their con-
tinued safety.
G. M. A.
Gordons 9 Red Colobus
COLOBUS BADIUS GORDONORUM (Matschie)
Piliocolobus gordonorum, Matschie, Sitzb. Ges. Naturf. Freunde Berlin, 1900,
p. 186. (Uzungwe Mountains, Uhehe, Tanganyika Territory.)
ORDEE PRIMATES: PRIMATES 163
This rather strikingly colored subspecies is known only from a
circumscribed area in the Uzungwe Mountains to the northeast of
Lake Tanganyika. The name was based on a single imperfect skin
found in a native hut and two other skins secured by the brothers
von Gordon, for whom it is named. In 1923 Kershaw recorded an-
other specimen secured in the same region by Arthur Loveridge,
who on a second visit obtained four others for the Museum of Com-
parative Zoology.
The top of the head is deep ferruginous, the back shining black;
forelimbs black, hind limbs mixed black and silvery, the tail mixed
black and ochraceous; lower surfaces white.
On the somewhat isolated Uzungwe Mountain range Mr. Loveridge
found this monkey but once, when at an altitude of some 5,000 feet
he came upon a troop just at dusk. They live in high forest and are
with difficulty obtained. Their nearest relative is perhaps Kirk's
Red Colobus of Zanzibar. Their chief danger is perhaps from native
hunters, but also possibly in future encroachments upon the small
area of forest to which they are confined.
G. M. A.
Kirk's Red Colobus
COLOBUS BADIUS KIRKII Gray
Colobus kirkii Gray, Proc. Zool. Soc. London 1868, p. 180, May, 1868. ("Zan-
zibar.")
FIGS.: Gray, op. cit., pi. 15; Elliot, 1913, vol. 3, pi. 16 (skull).
Restricted to the Island of Zanzibar, this form is in danger only
to the extent that future settlement and cultivation may reduce its
area of habitat.
Forehead and sides of head yellowish white, the long hairs extend-
ing beyond the sides of the head; crown, lower part of neck, back
from shoulders, reddish brown; shoulders, outer side of arms, hands
and feet black; throat and under parts grayish white; tail dull
reddish brown.
The chief interest of this monkey, apart from its island habitat
to which no member of the black-and-white group extends, is that
in the skull the median frontal suture remains open into adult life,
a rare condition sometimes found also in man.
This monkey was first secured and sent to Europe by Sir John
Kirk in 1868. He regarded it as rare at that time but in 1884 wrote
that it was still to be found in many of the wooded districts of the
island, although "so rare as not to be procurable, even when I sent
the hunters over the island. I have a report that it exists still in
one spot, which they could not reach. ... It looks as if the animal
will be lost to science. This is due to the destruction of forest and
jungle over the island." Two years later Sir Harry Johnston wrote
164 EXTINCT AND VANISHING MAMMALS
that it had "disappeared from nearly every part of the island of
Zanzibar, but a rumor prevailed that -it still lingered on a clump of
forest as yet unvisited by hunters." On sending his hunters thither,
they returned after a week's absence, bringing 12 dead monkeys,
with the report that they had killed every one, so that, as Sir Harry
supposed, this animal too had gone to "the limbo of species ex-
tinguished by the act of man." Nevertheless these evidently were
not the last, and even to this day a few still remain on the island,
but of their number and present status no information is at hand,
beyond the fact that Arthur Loveridge procured a pair there in 1923.
G. M. A.
Family PONGIDAE: Anthropoid Apes
The two forms of Gorillas (genus Gorilla) and the four forms of
Chimpanzees (genus Pan) are found in central Africa. The third
genus of the family, the Orang-utan (Pongo) , is represented by one
species, occurring in Borneo and in Sumatra. As man's nearest
living relatives, these apes have an exceptional interest for us, and
their generally waning numbers call for a discussion of each form
in this volume.
Orang-utan. "Mias" (Borneo) ; "Mawas" (Sumatra)
PONGO PYGMAEUS (Hoppius)
Simia pygmaeus Hoppius, Amoenit. Acad., 1763, p. 68. (Locality unknown.)
SYNONYMS: Simla satyrus Linnaeus, Syst. Nat., ed. 12, vol. 1, p. 34, 1766 (not
of the 10th ed.) ; Pongo wurmbi Tiedemann, Zool., p. 329, 1808 (Borneo) ;
Simia morio Owen, Proc. Zool. Soc. London 1836, p. 92, 1837 (Borneo) ;
Simla abelii Clarke, Asiatick Researches, vol. 16, p. 489, pis. 1, 3, 4. 5,
1825 (Sumatra). For extensive synonymy, including names given by
Selenka and others, see Elliot (1913, vol. 3, pp. 192-195).
FIGS.: Elliot, 1913, vol. 3, pis. 5 (photos of animal), 23-28 (skulls); Carpenter
and Coolidge, 1938, fig. opp. p. 18; Yerkes and Yerkes, 1929, figs. 43-66.
It is at present believed that the Bornean and Sumatran Orangs
are not separable even subspecifically ; at least they represent the
same species and are not found living elsewhere. No doubt they in-
habited the Asiatic mainland at no very distant time, but with the
separation of Borneo and Sumatra from the Malay Peninsula the
populations of these islands were cut off and have survived to the
present. On the mainland, remains of anthropoids resembling the
Orang are known from the Siwalik Hills of India, but there seems
to be no evidence of their survival to the historic period.
The adult Orang-utan is a large shaggy animal, of dark rufous
color. The profile of the skull is much more sloping than in the
African anthropoids, the skull showing very little of the brow ridges
ORDER PRIMATES: PRIMATES 165
so prominent in the latter. The arms are very long, reaching to
the ankles when the animal is erect; foot long and narrow, the
great toe very short. Tail absent. Prominent cheek callosities some-
times present in adult males. Wallace, who measured 17 freshly
killed Orangs, states that adult males "only varied from 4 feet 1
inch to 4 feet 2 inches in height, measured fairly to the heel, so as
to give the height of the animal if it stood perfectly erect; the extent
of the outstretched arms from 7 feet 2 inches to 7 feet 8 inches." The
total length of a Bornean skull is said to be 246 mm. (Elliot) , for
the largest of many. Selenka gives series of measurements. For an
excellent account of the history, characters, psychology, see Yerkes
and Yerkes (1929).
The Orang is a much more lethargic animal than the African
anthropoids, moving leisurely through the forest, seeking various
fruits, especially those of the durian, of which it is extremely
fond. Leaves and bark of certain trees are also eaten. It is some-
what social and may be found singly or in pairs or in small groups.
Banks (1931) writes:
In a wild state and unmolested, Mias exhibit little more than a benevolent
curiosity towards men and the extremely child-like and almost pathetic
expressions that can be assumed in captivity point to the Mias as an extremely
peaceful and gentle animal when left to himself, always remembering of
course that both temper and strength are there in reserve for use when
aroused. . . . [They] make a kind of platform of sticks on which they sleep
at night and even during the day but I have never seen captive ones make
any sort of roof or make use of leaves to keep the rain off, as is sometimes
alleged. Nests are of two kinds, either a flat platform or more usually a
deep triangular shaped affair in the upright fork of a tree. ... I counted
eleven such nests still with green leaves all close together near a "Kayu
Ara" fruit tree where a pair were feeding. . . .
The distribution of the Mias in Sarawak is peculiar in its relations to the
rest of Borneo; it occurs in parts of N. Borneo . . . and it is common in
W. Borneo, the Landak River and right up the Kapuas River. Now the
Mias is very sensibly fond of neither cold nor rain, in fact the damp is his
worst enemy and for this among other reasons the occurrence of Mias at 3000
ft. is very exceptional nor is he as common in the immediate lower vicinity
of mountains as he is at the foot. For some 70 miles the Kalinkang Moun-
tains run N.E. and S.W. forming a watershed between that part of the Kapuas
River running S.W. and numerous short Sarawak rivers running West into
the sea and it is obvious that these mountains form an obstacle to the
movements of Mias which are common on the Kapuas and curiously on
the Sarawak side. The explanation lies I think in a gap in the Kalinkang
Mountains which towards Lobok Antu slope away almost to sea level,
eventually to rise on the other side of the Batang Lupar Mts. and stretch
away unbroken northwards into central Borneo. It is therefore more or less
true that the Mias is confined to a range bounded on the N.E. by the Rejang
River, on the west by the Sadong River: the Orang Utan has flowed through
from Dutch Borneo and filled up suitable and available places.
In upper Sarawak the Orang was formerly reported, but these
reports are doubtful, although Everett records two imperfect skulls
166
EXTINCT AND VANISHING MAMMALS
found in a crevice of the limestone hills at Paku, but at present
it is absent from all that region adjacent to the watershed of the
Landak River. According to Wallace, it has a wide distribution
FIG. 19. Orang-utan (Pongo pygmaeus). From photo of specimen in
Philadelphia Zoo.
in the low country, inhabiting many districts not only on the south-
west and southeast, but also on the northeast and northwest coasts,
but of its more exact local distribution at the present time little
information is at hand.
In Sumatra, the only place outside of Borneo where the Orang
is native, it is now confined to the former state of Atjeh, which
ORDER PRIMATES: PRIMATES 167
comprises practically the northwestern quarter of the island. In
their recent report on the animal here, Carpenter and Coolidge
(1938) indicate that the regions of Lami, Tapa Toean, and Ba-
kongan are the places on the west coast of Atjeh where Orang-utans
live in the largest numbers. They occur over the greater part of
Atjeh except the high central districts above 1,500 meters, the cul-
tivated and thickly populated districts on the east coast, the grass-
lands of the north, the rough mountains north of Lamno, and culti-
vated sections of the west coast, especially around Meulaboh.
Although an estimate of the numbers and normal density of popu-
lation is largely guesswork, it seems likely that the centers of abun-
dance are "around the Simpang Kanan and Peureulak Rivers on
the East Coast, and along the West Coast in suitable forests from
Lami to Singkel." They show a marked preference for lowlands
but may range up to 1,500 meters, though with a sharp decrease
above 700 or 800 meters. There is some evidence of local move-
ments following the seasonal ripening of certain fruits on which they
feed. In summary, these authors state that the Orang is found in
"an estimated fifty per cent of the primary forests" in Atjeh; "as
large clearings are made in* the rather level lowlands, these apes
are being destroyed or forced into the hills and mountains where
it is questionable whether or not conditions, including food supply,
are sufficiently suitable for the maintenance of the present popula-
tion level. However, large areas of Atjeh, because of its rugged
topography and inaccessibility, will remain naturally protected for
a long time as an orang-utan habitat. Europeans and not natives
threaten the orang-utan population, the most serious inroads being
made by commercial developments in the areas suitable for orang-
utan habitats. Numbers of these apes are being shot annually under
the supposition that they attack human beings, and it is feared
that government records do not accurately record all animals cap-
tured or killed."
For the better preservation of these interesting apes, the authors
recommend an extension of the present Alas National Park or Loser
Reservation and the development of a smaller reserve especially for
Orangs south of Meulaboh, together with the planting of food trees,
especially the durian. Further, the desirability of additional reserves
and special measures is advocated, particularly that "the killing
and capture of this animal for trading or exhibition purposes be
completely stopped and that its use for accredited but limited scien-
tific purposes alone be permitted."
G. M. A.
168 EXTINCT AND VANISHING MAMMALS
Coast Gorilla. Gorille (Fr.). Gorilla (Ger.)
GORILLA GORILLA GORILLA (Savage and Wyman)
Troglodytes'] gorilla Savage and Wyman, Proc. Boston Soc. Nat. Hist., vol. 2,
p. 245, 1847. ("Empongwe, near the river Gaboon, Africa"; about lat.
20' N., long. 9 30' E.)
FIGS.: Gervais, Hist. Nat. Mammiferes, pt. 1, pis. facing pp. 26, 28, 1854;
Du Chaillu, 1861, frontisp.; Forbes, 1894, vol. 2, pi. 28; Proc. Zool. Soc.
London 1896, p. 505, fig.; Elliot, 1913, vol. 3, frontisp.; Cunningham,
1921, pp. 119-124, figs.; Barns, 1923, figs. 45, 46, 50, 51; Yerkes and Yerkes,
1929, numerous figs.; Coolidge, 1936, pi. 12; Raven, 1936a, p. 316, fig.;
Fauna [Philadelphia], vol. 1, no. 1, pp. 8-9, figs., 1939.
The Coast Gorilla is generally considered to be diminishing in
numbers but not to be in danger of extermination. All gorillas are
given full protection under Schedule A of the London Convention
of 1933.
"This animal ... is much larger and more ferocious than the
Chimpanzee. Its height is above five feet; but it is remarkable for
the disproportionate breadth of the shoulders, which is double that
of the Chimpanzee. The hair is coarse, and black, except in old
individuals, when it becomes gray. The head is longer than that of
an ordinary man by two inches, and is remarkable for having a crest
of coarse hair over the sagittal suture, which meets at right angles
a second, extending over the upper part of the occiput, from one
ear to the other. The fore-arm is much shorter than the arm, the
hand is remarkable for its great size, and the thumbs larger than
the fingers." (Savage and Wyman, 1847, pp. 245-246.)
"Face and chest bare, black; . . . arms and belly black; back
and outside of thighs gray grading into black towards ankles and
on feet; hands black; no beard; top of head black, nape mixed
black and red." Height, 5 feet 10 inches. (Elliot, 1913, vol. 3,
pp. 213-214.)
The numerous described forms of Gorillas are reduced by Cool-
idge (1929, p. 348) to two subspecies the present one and the
Mountain Gorilla (G. g. beringei).
"For the Coast Gorilla, the westernmost boundary approximates
the Cross River in the southern provinces of Nigeria. The most
westerly point actually recorded is Ikom, 8 40' east and 6 north.
The northernmost point is close to Basho, 9 25' east, 6 7' north.
On the east we have reports from several places such as Wesso and
Nola on the Sanaga [error for Sanga] River. The Sanaga River,
about 16 15', seems to mark the eastern boundary of the range
of the Coast Gorilla. On the southeast the line follows the border
of the forest which reaches its southernmost limit at Mayombe on
the edge of the Belgian Congo, 5 south, 13' [ = 13] east. Along
the Atlantic coast in most places the forest begins a little way inland.
ORDER PRIMATES: PRIMATES 169
Gorillas have been reported actually on the coast, but generally they
are found not closer than thirty miles from the sea. They seem
especially plentiful along the Gaboon, Ogowe, Camp, and Sanaga
Rivers." (Coolidge, 1929, p. 363.) More recently the range has been
found to extend somewhat eastward of the Sanga River in French
Equatorial Africa (Coolidge, 1936, p. 493, maps 1-2). The total
range of the Coast Gorilla seems to be strictly confined to the west-
ern portion of the Lower Guinea Forest District of Chapin (1932,
p. 90) and of Bowen (1933, pp. 256, 258).
Gabun. "They live in herds, the females exceeding the males in
number. . . . They are exceedingly ferocious, and objects of terror
to the natives, who seldom encounter them except on the defensive.
The killing of an Engeena is considered an act of great skill and
courage, and brings to the victor signal honor. . . .
"Their flesh, when obtained, is eaten by the natives, as well as
that of the Chimpanzee." (Savage and Wyman, 1847, p. 246.)
Du Chaillu's classical account of the Gorilla (1861, pp. 388-
404) is based upon his experiences in Gabun. Among other things,
he says (pp. 399-400) : "The negroes never attack them with other
weapons than guns; and in those parts of the far interior where no
European guns had yet reached, as among the Apingi, this great
beast roamed unmolested, the monarch of the forest."
"The Fernan Vaz District ... is considered the best region
for both the gorilla and the chimpanzee." One "family party of
five or more gorillas" was encountered, and on another occasion
"a large family" was reported. The animal will occasionally ad-
vance to attack when not wounded. (Aschemeier, 1921, pp. 90-92.)
"The majority of the Gaboon skulls have come out from the
region of the Ogowe River" (Coolidge, 1929, p. 303).
A. R. Maclatchy (in litt., February 5, 1937) gives the following
report for Gabun: "The decree of 1929 classed the gorilla among
the protected species. Its great vulnerability makes it an adversary
much less dangerous than the buffalo and the elephant. It rarely
pushes its attack to the limit. Sometimes it visits the native plan-
tations by night. Its food consists of various plants. I do not see
the reasons for a protection as strict as that which the gorilla
enjoys. It is little hunted, except in legitimate defense, by the
natives, who have a superstitious terror of it, and by a few hardy
European sportsmen. It scarcely seems to be threatened with de-
struction. It really abounds, and is protected by its habitat and
by its natural shyness. More specimens could be allowed on hunting
permits without the risk of diminishing its numbers."
Cameroons. In the southeastern corner of Cameroons, and per-
haps also across the boundary in French Equatorial Africa, the
Gorilla appears quite common, and almost entirely inoffensive as
170 EXTINCT AND VANISHING MAMMALS
to attacks on man. The forest is so dense, and the animal so shy,
that it is extremely difficult for the hunter to get sight of it. Thus
the Gorilla seems fairly well protected. Bands numbering up to more
than 20 individuals are reported. (Ramecourt, 1936, pp. 217-247.)
Raven (1936a, 19366) gives a most interesting and detailed
account of hunting Gorillas during more than a year spent at va-
rious places in southern Cameroons. The natives here are very keen
to eat Gorilla meat, being generally faced with a deficiency of
meat in their diet. A missionary reported many of the animals at
Djaposten, in southeastern Cameroons, where "in one morning's
walk of perhaps two hours he had counted more than 100 gorilla
beds." Although Raven himself found the animals quite common in
this region, it was extremely difficult to obtain a good view of them
in the dense forest, and only three adults were collected during his
entire sojourn, despite assiduous hunting. Raven writes further
(19366, pp. 529-530) :
For centuries past the gorillas and natives have been competitors. As the
native populuation increased, new villages would be formed and more
clearings made. Then epidemics would occur, killing off great numbers of
natives, and their gardens would be neglected to run into second growth.
The gorillas, with a constitution so nearly like that of man that they can
find more food in human plantations than in the virgin forest, would move
into these deserted clearings. There with an abundance of food they throve
and congregated, to such an extent eventually that if only a few natives
remained they were actually driven out because of their inability to protect
their crops against the gorillas. But with the advent of the white men's
government, with the distribution of firearms among the natives, preventive
medicine and the treatment for epidemic and infective diseases, man has the
upper hand at present in this age-long struggle.
"Mr. Raven had opportunity to witness the unfortunate effect,
so far as the protection of the gorilla was concerned, of the demand
for gorilla skulls on the part of scientists, to such a degree that
white men as well as natives had in the past often done a profitable
business in killing the animals and selling their skulls. The result
had been a rapid decrease in the gorilla population, so that Mr.
Raven, although by his record known to be a hunter and collector
of the first rank, was compelled to hunt week after week in a des-
perate effort to come up with the nervous survivors of the race in
this district. . . .
"Mr. Raven's experience leads him to believe that ... the gorilla
is being rapidly exterminated in many localities." (Gregory, in
Raven, 19366, p. 540.)
It is doubtful if the protective laws have stopped the killing of
Gorillas by natives to any extent. Most of the museum specimens of
skulls, etc., are from native-killed animals and have been turned
ORDER PRIMATES: PRIMATES 171
in to traders. Natives capture them by spearing and by snaring.
(H. C. Raven, oral communication, March 17, 1937.)
The Gorilla is utilized for experimental purposes in the study of
human diseases (Ministry of Colonies, Paris, in litt., November 7,
1936).
Gorillas are decreasing but not disappearing. The cause of
depletion is native hunting for food. (Inspection of Waters and
Forests, Yaounde, in litt., January 12, 1937.)
The number in French Cameroons is estimated at some thousands.
They are partially protected by law, one head being allowed on a
full license. (Paris Agency, in litt., November, 1936.)
Nigeria. Coolidge (1929, p. 303) refers to the range as including
"the Western Cameroons [part of Nigeria], which is a comparatively
limited section centering around Mamfe or Dakbe and extending
west as far as the Cross River. A great many skulls come from this
region."
Hay wood (1932, p. 32) reports the species from the borders of
Ogoja and Cameroons Provinces.
In British Cameroons, Sanderson (1935, p. 26) reports Gorillas
from the mountains of Assumbo, about the headwaters of the Cross
River. They "are numerous in the Mountain Moss Forest belt,
where the natives record their movements minutely."
"In Nigeria where a few exist the natives take an annual toll and
I do not think there are many" (C. W. Hobley, in litt., August 18,
1936).
French Equatorial Africa. The Gorilla does some damage in the
banana plantations, but it is not important. It seldom attacks man
without provocation. There is no reason why it should not be abso-
lutely protected. Its northern and western limits are unknown; its
southern seems to be the Congo. (Lavauden, 1933, p. 30.)
"Four complete specimens of the Coast Gorilla were procured
by the Vanderbilt Expedition of the Academy of Naturaf Sciences
of Philadelphia in the winter of 1934." Three "were killed by natives
in the neighborhood of Aboghi, forty miles southwest of Nola near
the west bank of the Sanga River." The fourth was secured "near
Barundu, about 15 miles east of the Sanga River and 22 miles north-
east of Nola." (Coolidge, 1936, p. 479.)
Green (in Coolidge, 1936, pp. 491-492) reports Gorillas as abun-
dant in the region of Aboghi. "The old males appear to be somewhat
solitary, but small bands of four to ten were noted from tracking."
"The expedition reported that gorillas were frequently killed on
the left bank [of the Sanga]. . . .
"The field notes of Mr. Rehn and Mr. Green give us the impression
that gorillas were plentiful in the region from which these specimens
172 EXTINCT AND VANISHING MAMMALS
came, and that they are frequently hunted by the natives." (Cool-
idge, 1936, pp. 493, 499.)
According to all accounts, Gorillas still occur in fair numbers in
the Sanga River region, though not so commonly as 25 years ago.~A
local French doctor, in the course of two years, had treated nine
natives for Gorilla attack, one of the cases being fatal. (J. A. G.
Rehn, oral communication, March 22, 1939.)
Belgian Congo. Schouteden (19306, pp. 298-299) presents evi-
dence of the rare occurrence of Gorillas in the Mayumbe forest,
north of the lower reaches of the Congo River. Later (19366,
pp. 15-16) he records a skull from the Haut Mayumbe. Here the
animal had seemed to have disappeared, or to occur only occasion-
ally, coming perhaps from Gabun. But it appears to occur still in
certain parts of Mayumbe, thanks, perhaps, to the protection it has
enjoyed for some years.
Use in research. Yerkes and Yerkes point out (1929, p. 590)
that the Gorilla and other anthropoid apes "must inevitably become
the preferred substitutes for human subjects in investigations which
may not be carried on with the latter and which have as objectives
the extension of knowledge and control of human life." They also
stress (p. 589) the greater availability and controllability of these
animals for use in the investigation of various problems in genetics,
physiology, neuro- and psychopathology, psychology, sociology,
pedagogy, and experimental education.
Survival Yerkes and Yerkes (1929, p. 396) quote Keith (1896)
as follows: "From accounts furnished by travellers and hunters, one
infers that the total population [of the species as a whole] males,
females, and young is well under 10,000." Eventually (1914)
Keith raised this estimate to 20,000 to 30,000 individuals, but
Yerkes and Yerkes remark (p. 397) that it is difficult to decide how
seriously this estimate should be taken. They also say (p. 397) :
"Concerning abundance or frequency little is known. Both early
and late in the last century the relative rarity of the gorilla sug-
gested to investigators its disappearance and probable extinction.
From limited distribution, difficulty of negro hunters in procuring
skins of adults, and the small number of captive specimens sent to
Europe, Deniker (1891, pp. 369-370) infers that the process of
extinction is under way. . . .
"For nearly a century it has been known that the gorilla is the
rarest of the manlike apes."
H. C. Raven (oral communication, March 17, 1937) estimates
the total number of all Gorillas now living at more than 1,000 and
at less than 10,000.
ORDER PRIMATES: PRIMATES 173
Mountain Gorilla. Gorille ties montagnes (Fr.)
GORILLA GORILLA BERINGEI Matschie
Gorilla beringeri [misprint for beringei] Matschie, Sitz.-Ber. Ges. Naturf.
Freunde Berlin 1903, no. 6, p. 257, 1903. ("Auf der Spitze des Vulkans
Kirunga ya Sabinyo in einer Hohe von 3000 m," German East Africa;
i. e., Mount Sabinio or Sebyinyo, at the boundary point of Ruanda,
Uganda, and the Belgian Congo. Not at the summit, but on the south
or southeast flank, at about 2800 m. (Derscheid, 1928, p. 150).)
FIGS.: Lonnberg, 1917, pi. 1; Barns, 1922, frontisp. and pis. facing pp. xvi,
83, 86; Barns, 1923, figs. 43, 44, 52; Akeley, 1923a, pp. 428, 438, 440,
444, figs.; Akeley, 19236, frontisp. and pis. facing pp. 190, 206, 222, 230;
Yerkes and Yerkes, 1929, numerous figs.; Coolidge, 1929, pi. 1, and 1930,
pp. 626-627, figs. 454, 454b; Raven, 1931, cover and p. 241, fig.; Bingham,
1932, pis. 18, 19, 22; Jour. Soc. Preservation Fauna Empire, n. s., pt. 18,
frontisp., 1933.
The Mountain Gorilla is now well protected in its range centering
in the Pare National Albert in the eastern Belgian Congo, and its
chances for survival appear to be excellent.
Face, ears, breast, back, hands, and feet naked; breast brownish,
like worn leather; back somewhat lighter; face, ears, and naked
parts of the limbs black; hair black, long, and thick, and forming a
pronounced beard on cheeks and chin. Height 1.5 m.; weight 100 kg.
(Matschie, 1903a, p. 254.) "The external characters that distinguish
the Mountain from the Coast Gorilla are, besides a longer palate
and a generally narrower skull, the thicker pelage, shorter arms
and longer legs, large amount of black hair, and fleshy callosity on
the crest" (Coolidge, 1929, p. 375). "The large patch of silver-gray
fur covering the back of the adult male gorilla is the most remark-
able part of his coloration; the female is entirely black, and very
much smaller than her mate" (Barns, 1923, p. 130) .
Coolidge (1929, p. 363) says of its range:
The Mountain Gorilla is found in a comparatively narrow strip of the
eastern Congo. Its principal habitat is the mountain forest as distinguished
from the lowland forest of the Belgian Congo. Its northern limit is Mulu,
10' south, 29 10' east (Absil and Chapin). We find it as far west as
Walikale, 1 20' south, 28 1' east, where it strays a little into the lowland
forest. The eastern limit seems to be close to Kigezi in Uganda, 1 15' south,
29 45' east. The southern limit is Baraka on Lake Tanganyika, 4 19' south,
29 2' east. In this entire region the gorillas that are most known and accessible
are the troops that inhabit the volcano regions where Akeley died while
studying them. Whether they are entirely isolated from contact with outside
gorillas at the present time is doubtful and has not yet been established.
In the mountains back of Baraka, Boko, Uvira, and Katana large troops
have been recently found in the upland forests.
"I have examined . . . the sources of evidence for the existence
of gorillas in the intervening area between (longitude 17 east) the
eastern limit of the known range of the Coast Gorillas and (longi-
174 EXTINCT AND VANISHING MAMMALS
tude 28 east) the western limit of the known range of the Mountain
Gorillas. With a single exception, I attach no great importance to
this evidence. The exception refers to the four skulls from Bondo
on the Uelle River collected by Lemarinel in 1908. These furnish
us with definite proof for the existence of gorillas in the Djabbir
region as late as 1908. Except for these skulls no other tangible
evidence of gorillas in a forest belt of 650 [ = about 750] miles has
turned up." (Coolidge, 1936, p. 500.) Coolidge considers (p. 497)
that the affinities of the Bondo skulls are with the Coastal Gorilla,
but G. M. Allen (1939, p. 177) refers G. uellensis Schouteden, which
was based upon these skulls, to the synonymy of beringei.
The Mountain Gorilla was first made known to science through a
specimen shot by Capt. Oscar von Beringe on Mount Sabinio about
1902 or 1903 (Matschie, 1903a, p. 253).
In 1913 and 1914 seven specimens were obtained by E. Arrhenius
on the volcano Mikeno, Virunga Mountains. "According to Captain
E. Arrhenius the Gorillas are rather numerous .... They live in
bands consisting of 20-30 individuals ....
"The natives hunt the Gorillas to obtain their skin which they
use for wrapping up their copper thread etc., or for revenging some
relative. Thus when a man from Sangana had been killed by a
Gorilla his family killed five Gorillas in revenge. The natives hunt
Gorillas with the aid of dogs. The dog bites the Gorilla and returns
to his master who waits for the Gorilla with the spear ready. He
throws the spear at the Gorilla and runs away. The dog repeats the
maneuvre, until the animal is killed. The natives do not eat the
meat of Gorillas, nor that of Chimpanzees." (Lonnberg, 1917, pp. 7,
17-18.)
Barns (1922, pp. 81-88) encountered a band of Gorillas between
the volcanoes Mikeno and Karisimbi, and secured a specimen, which
his hungry native porters refused to eat. "This monster ape would
seem to have no enemies, failing man; and even man, the most
dreaded of all the animal world, holds little fear for the gorilla in
his inaccessible home" (p. 87) .
"Its food consists, apart from bamboo shoots, entirely of herbage
docks, sorrels, hemlocks, etc. although honey may be part of the
menu. He does not grub for roots, neither does he eat fruit as a
general rule ....
"Savage man, through superstition as much as anything else,
but also on account of the inaccessibility of the gorilla's mountain
home, has left this ape unmolested; we therefore find him and his
family habitually and fearlessly sleeping on the ground." (Barns,
1923, pp. 129-130.)
"As regards longevity, gorillas, on account of their life free from
molestation, famine, or disease, and also judging by the worn teeth
ORDER PRIMATES: PRIMATES 175
of one animal I secured, live, in my opinion, to be a much greater
age than man" (Barns, 1923, p. 132).
"The natives of this region have no fear of the gorilla. . . . Some
of my guides and my gun bearer were trappers and hunters in the
gorilla forests and were thoroughly familiar with them. At no time
did the guides or gun boys show any indication of anything more
than casual interest even when we approached very close to
gorillas." (Akeley, 1923a, pp. 438-439.) Akeley continues (p. 447) :
After my first expedition into the gorilla country, I am more convinced
than ever not only that the gorilla is one of the most fascinating and impor-
tant objects of study in the realm of natural history, but also that his dis-
position is such as to permit the most intimate observation of his habits.
... A few weeks of casual acquaintance and one is fired with a desire to
ferret out the answers to a hundred questions about this little-known relative
of man questions of increasing importance to scientists and physicians in
their efforts to understand and aid man himself. Probably no other project
of so moderate a size is likely to lead to such immediate and valuable scien-
tific results as that which will make of the Kivu region a sanctuary, where
the gorillas under the protection of man may grow more and more accustomed
to human beings and where through a series of years they may be observed
and studied.
On the three mountains, Mikeno, Karisimbi, and Visoke, "I judge
that there are between fifty and one hundred animals altogether"
(Akeley, 1923b ; p. 248).
Akeley's efforts led to the establishment of the Pare National
Albert, comprising the Kivu volcanoes and providing for the special
protection of the Mountain Gorillas. Meanwhile Burbridge had
estimated their numbers at 1,000 to 2,000 individuals. Their range
extends beyond the volcano region to the bamboo forests dominating
the highlands of the Great Lakes. (Leplae, 1925, pp. 15, 19.)
According to Derscheid (1928, pp. 154-159), the animals are
especially numerous at elevations between 2,700 and 3,500 m., with
extreme occurrences at 1,900 and 3,900 m. He has met with a few
solitary old males, but more usually with bands of 7 to 43 indi-
viduals. He estimates the number on the central massif (Mikeno-
Karisimbi-Bishoke) at 350 to 500; on the eastern massif (southern
slopes of Muhabura, Sebyinyo, and Mugahinga) at 150 to 200; and
in the Uganda portion of the region (northern slopes of the three
volcanoes just mentioned) at 100 to 150. He also remarks on the
surprisingly small proportion of young animals among the Gorillas
observed.
"During our two months stay among the different peaks of the
Birunga Range we observed several herds of Gorillas. The largest
of these herds consisted of about 20-30 individuals .... In all
about 70 examples were seen by the members of the Expedition.
. . . Their stronghold seems ... to be the mountain triangle com-
posed of Mikeno, Karissimbi and Vissoke. They are mostly found
176 EXTINCT AND VANISHING MAMMALS
in the Bamboo Region, but they also live higher up the steep moun-
tain slopes with their beautiful vegetation of Hagenia-trees." (Gyl-
denstolpe, 1928, p. 23.)
"There is a spotted menace, a potent factor too, in the leopard,
who destroys numbers of young animals" (Burbridge, 1928, as
quoted in Yerkes and Yerkes, 1929, p. 398) .
Pitman (1935, pp. 477-494) gives an excellent account of Gorillas
in Uganda, and the following excerpts are taken from his paper:
The occurrence of Gorillas in the Kayonsa region of Uganda [about midway
between the Birunga volcanoes and Lake Edward] has been known for many
years (p. 477).
There is in the Kayonsa a complete absence of bamboo, wild celery, dock,
and similar juicy-stemmed plants such as abound in the humid, high alti-
tudes, forcing the Gorilla to confine its diet to a mixture of leaves, berries,
ferns, the tender fronds of tree-ferns, parts of the wild banana stems, and
leaves, and fibrous bark peeled off a variety of shrubs in the undergrowth. . . .
Owing to a lack of what apparently are normal food constituents the
Gorilla has become more enterprising in search of food, and in consequence
climbs trees freely to a known height of at least 50 feet. (P. 478.)
The "beds" of the Kayonsa Gorilla are large platforms built in the trees,
and often at a considerable height above the ground.
[The altitude of the habitat varies between 6000 and 7900 feet.] (P. 479.)
The forest region to the east of the Kishasha river [where some Gorillas
are known to occur] is a gazetted forest reserve and, in consequence, not
open for human settlement. There is little likelihood in the immediate
future of serious conflict between Man and Gorilla in the dense uninhabitable
valleys to the west of this river and in the vicinity of the Belgian Congo
border ....
It was calculated that this western area harboured forty to fifty Gorillas.
[In the entire region there were possibly at least eighty.]
Normally the troops vary in size from five to eight or nine, [but one troop
was said to include nearly two dozen]. (P. 480.)
The [Kayonsa] Gorilla normally is peaceably disposed and not aggressive
(p. 483).
The Wambutte [Pygmies] are extremely tolerant of the Gorillas, but not
so the other local natives, who would readily endeavour to exterminate the
lot, were it not for the fact, of which they are well aware, that these splendid
animals are absolutely protected (p. 484).
The animals are said to sometimes raid the native gardens but
not to attack the natives.
The Chimpanzees
In spite of the multiplicity of names that have been applied to
the Chimpanzees, it seems probable that only four valid forms are
recognizable, representing probably two distinct species, as follows:
Common Chimpanzee
PAN TROGLODYTES TROGLODYTES (Blumenbach)
Simla troglodytes Blumenbach, Handb. der Naturgesch., p. 65, 1799. ("An-
gola.")
ORDER PRIMATES: PRIMATES 177
Long-haired Chimpanzee; Eastern Chimpanzee
PAN TROGLODYTES SCHWBINFURTHII (Giglioli)
Troglodytes schweinjurthii Giglioli, Ann. Mus. Civ. Stor. Nat. Geneva, vol. 3,
p. 114, footnote, 1872. (Upper Uele drainage, Niam-niam country, eastern
Congo Beige.)
Western Chimpanzee
PAN TROGLODYTES VERUS Schwarz
Pan satyrus vents Schwarz, Ann. Mag. Nat. Hist., ser. 10, vol. 13, p. 578,
June, 1934. ("Sanda Magbolonto chiefdom, Karima district, Sierra Leone.")
SYNONYMS: For list of synonyms of these three races, see G. M. Allen (1939b,
pp. 172-175).
FIGS.: Elliot, 1913, vol. 3, pis. 7, 8, 8 bis (animal); pis. 36-39 (skulls); Yerkes
and Yerkes, 1929, figs. 69-118.
Yerkes and Yerkes (1929) write that the "description of the con-
figuration of the type chimpanzee is as difficult as description of
man, so numerous and pronounced are individual, sex, and species
differences and developmental changes." In general, of anthropoid
form, the forelimbs proportionately long, reaching below the knee
when the animal stands erect. Form stocky, shape of ear much as in
Homo, forehead heavily ridged, nose flattened. Face usually bare
or nearly so, and in the adult black like the skin of the body, except
in the race verus, in which it is paler. Hair of the head directed
backward in the typical race, but usually with a parting in verus.
In the eastern race, schweinjurthii, the hair is longer than in the
others. The maximum (standing) height of the male is about 5
feet, of the female 4 feet. Weight of male 125 to 175 pounds; of
female 100 to 150 pounds. The skull is distinguished readily from
that of a Gorilla by the smaller teeth and by the fact that when
viewed from in front the summit of the brain case is visible above
the brow ridges instead of being hidden by them. Color usually
black, with often a whitish pygal patch.
Throughout the vast extent of the tropical rain forest from the
Gambia and adjacent French Equatorial Africa, south to the
Congo, and eastward to the borders of Uganda and Tanganyika,
Chimpanzees are found, but they vary greatly in local abundance.
They seem much given to wandering about over circumscribed
areas, and so it is difficult to make censuses or to estimate popula-
tions. Moreover, the nature of their habitat in rain forest of dense
growth makes their observation uncertain. Thus in our journey
across Liberia in 1926, a country in which they are believed to be
rather common, I saw none, and H. J. Coolidge, Jr., came upon them
but once in the eastern border of the country. Yet they are com-
mon in the region about Kindia, in French Guinea, and occur in
7
178 EXTINCT AND VANISHING MAMMALS
numbers in Sierra Leone and in the forests of the Belgian Congo and
in the Cameroons. The original specimen was said to have come
from "Angola" but probably was not native there, for the larger
species is not now known from south of the Congo. If it was actually
brought from there, it was no doubt purchased of natives who had
captured it as a young animal farther north.
The Chimpanzee offers no trophy for the sportsman and should
not be killed or captured except for scientific purposes. It is thus
included in Schedule B of the London Convention of 1933. Its natural
enemies must be few and, except for man, probably include only the
Leopard. Native peoples seldom molest them, except where there is
inducement from whites to capture them for "pets" or to secure
specimens. Many tribes believe that "every chimpanzee is linked
with the soul of a man, so that if one is killed the man too will die,"
or some other calamity will ensue. At Kindia in French Guinea, the
Pasteur Institute maintains a laboratory for the observation and
medical study of these animals, where individuals may be accus-
tomed to captivity before being sent to institutions in Europe or
elsewhere. "From the medical point of view, we have no need of
emphasizing the advantage to be derived to-day from anthropoids,
and especially the Chimpanzee, in the study of human diseases ; the
experimental inoculations of serums, vaccines, and medications of
all sorts, find in the Chimpanzee a very valuable subject" (Lavau-
den, 1933, pp. 30-31). Psychological studies of this animal have
already thrown much light on the evolution of intelligent behavior;
for a review of such work the reader is referred to the volume by
Yerkes and Yerkes (1929).
While the reports of comparative abundance, as noted by travelers
or persons stationed in parts of its range, are of only relative value
as often recording mere casual impressions, nevertheless the following
notes are here added as providing a brief survey of its occurrence
in selected stations. In the Gambia it is said no longer to exist near
the coastal towns, but according to E. Johnson (1937, p. 62) every
year "about fifty animals are brought in for sale from Futa Jalon,
some 70 miles southeast of Fatoto, 280 miles from Bathurst." They
are found in the Gola Forest Reserve of Sierra Leone, and small
troops may be met with by good fortune in the great forests of
Liberia. On the Gold Coast, according to Haywood (1933), "they
are only reported from the Western Forest belt, but it seems quite
possible they are spread over a large area, although by no means
in large numbers." The Director of Agriculture of the Gold Coast
writes (in litt., 1937) that the "chimpanzee is now rare and confined
to the extreme western border of the forest country, but whether
it was ever plentiful is not known." In Nigeria it is reported from
forest regions of Oyo, Onitaha, Owerri, Ijebu Ode, and Abeokuta
ORDER PRIMATES: PRIMATES 179
Provinces, so probably is present in Benin, Ondo, Calabar, and Warri
Provinces (Haywood, 1932) . It is apparently common in the Gabun,
and in the southern Cameroons. It is "numerous" in the equatorial
forests of the Belgian Congo but in the Ubangi-Shari district is found
only in the Ubangi Basin, in small numbers (at most a few hun-
dreds), localized in Haute-Sangha, Lobaye, Ouaka, and Haut-
Mbomou. It does not seem to have diminished except in the Ouaka.
In this region it was completely protected since 1916, then partly in
1931, and once more completely in 1936. It occurs also in the Pare
National d'Odzala in the Middle Congo and in general appears to be
threatened not with extermination but with diminution (L. Blancou,
in letter of 1937) . A. J. Jobaert, in response to queries, writes that in
the Belgian Congo very few are now killed by Europeans, "but cer-
tainly the natives, and especially the pygmies, destroy a considerable
number, although it is totally protected by law ; they were certainly
quite numerous a few years ago."
In the eastern part of its range, the race schweinfurthii is locally
common as far as the border of the rain-forest area in Uganda and
extreme western Tanganyika. In the upper Congo region, Lang and
Chapin found it common, as about Aba and Faradje on the north-
eastern border of the rain forest, and at Avakubi, Niapu, and Medje
within the forest. On the other hand, reports from the Uganda
Game Department (1928) indicate that in Kigezi "it would appear
that the numbers of this species are diminishing. It is, however,
likely that a permanent change of quarters has resulted in its dis-
appearance from localities where it was previously known. It has
been ascertained from Ankole that the parties or families of Chim-
panzees are great wanderers and not confined to specified localities.
However, information both from Toro and Bunyoro districts also
records a recession from areas in which till recently these animals
were seen and heard." In Tanganyika, Chimpanzees have been
recorded as far south as the eastern shore of Lake Tanganyika,
south of Kigoma, in the Mahare Mountains. Footprints and about
a dozen sleeping platforms were noted by B. W. Savory. They have
also been found on the west side of the lake as far south as the
Marungu district. Mr. Savory found collecting of Chimpanzees very
difficult here, not only on account of the nature of the country but
also on account of the superstitious fear of the natives, who believe
these apes are reincarnations of human beings and that a gun fired
at one of them will surely burst (Dollman, 19356, pp. 15-16) . On
Mount Kivu Chimpanzees are said to be found but are extremely
localized. Derscheid records them from Mount Henu and in the
bamboo forest south of the Karissimbi Volcanoes.
While such areas of rain forest as are found on some of the more
eastern isolated mountains, as Kilimanjaro, Kenya, and Elgon,
180 EXTINCT AND VANISHING MAMMALS
might apparently be suitable for Chimpanzees, and in some future
time might serve as sanctuaries for transplanted stock, they are at
present uninhabited by these apes. In the distant past, however,
they may have sheltered them. There is much evidence that in
eastern Africa the lowlands and slopes of mountains were anciently
clad with heavy forest but that native races of man have in the
course of centuries gradually, by burning and cutting, beaten back
this forest, and that it has given place to scrubby growth or finally
to scattered thorn scrub. A similar process is slowly going on both
within the rain forest and at its edges, with slow but gradual clearing
of trees for agriculture, and subsequent abandonment. Continued
long enough, this results in final destruction of the high forest, first
in spots, then in local areas, and finally over larger tracts, all of
which will eventually much curtail the available living areas for
Chimpanzees.
G. M. A.
Lesser Chimpanzee; Pygmy Chimpanzee
PAN PANISCUS Schwarz
Pan satyrus paniscus Schwarz, Revue Zool. Africaine, vol. 16, p. 425, April 1,
1929. (South of the upper Maringa River, 30 km. south of Befale, south
bank of the Congo, Congo Beige.)
FIGS.: Coolidge, 1933, pi. 1, figs. A, B; pi. 2, fig. A.
Present evidence seems to indicate that this is a smaller species
than the Common Chimpanzee and distinct from it. Its known
range is in the Middle Congo forests, on the south side of the river,
here supposed to form a physical barrier to northward extension.
Coolidge (1933) has summed up our knowledge of this animal and
has made a comparative study of its skeleton. An adult female
(containing a fetus) had a head and body length of 630 mm.;
height from crown to sole, 1,010; spread of arms, 1,510. It is thus
much smaller than the other species. The hair is fine in texture and
glossy black throughout except for a small white pygal tuft, and is
long and dense, without a parting on the head. The skull has a
rather juvenile appearance in its inflated forehead and small brow
ridges.
Although the existence of a Chimpanzee on the south side of the
Congo had been several times reported, it was not until 1928 that a
M. Ghesquiere obtained specimens for the Congo Museum in Ter-
vueren and the animal was described. Previously a specimen had
been in the British Museum, collected in 1895. Other specimens
have since been secured, and more information is likely soon to be
placed on record. Dr. James P. Chapin, who secured a specimen in
1930 near Lukolela, describes the voice as neither so loud nor so shrill
ORDER EDENTATA: EDENTATES 181
as that of the larger Chimpanzee. Dr. R. M. Yerkes had a specimen
in captivity for over a year in 1923-24 and has written of its
behavior. Very little is known of its abundance, but one may sup-
pose its distribution includes the rain-forest area between the
Congo and the Kasai.
G. M. A.
Order EDENTATA: Edentates
Family MANIDAE: Pangolins
Three genera of this family, represented by four species, occur
in Africa south of the Sahara, and all of them are treated in the
following pages. Two other genera (Manis and Phatages), repre-
sented by five forms, occur in the Oriental region (India, China,
Siam, Indo-China, Malaysia) ; while subjected to some perse-
cution by reason of their supposed medicinal value, these Oriental
pangolins are not included in the present report.
Giant Pangolin or Scaly Anteater. Pangolin geant (Fr.).
Riesenschuppentier (Ger.)
SMUTSIA GIGANTEA Illiger
Manis gigantea Illiger, Abhandl. K. Akad. Wissen. Berlin, physik. Kl., 1804-
1811, p. 84, 1815. (Guinea = West Africa.)
FIGS.: Buttikofer, 1890, vol. 2, p. 394, fig.; Beddard, 1902, p. 190, fig. 109;
Schubotz, 1912, p. 357, fig.; Bequaert, 1922, pi. 24, fig. 2, pi. 25, fig. 2;
Schouteden, 1930, p. [14], fig. 3a; Halt, 19346, pis. 32-34, and 1934c,
p. 727, upper fig., p. 729, fig.; Rosevear, 1937, p. 12, fig. 2.
The various species of African pangolins do not appear to be
numerous anywhere. They are in considerable demand among the
natives, and active hunting keeps down their numbers. All forms
are placed in Schedule B under the London Convention of 1933.
In all African pangolins "no hairs project between the scales, the
median dorsal row of scales does not extend to the tail tip, and
there is no external pinna of the ear." In S. gigantea and S. tem-
minckii "the belly is naked, the preaxial surface of the fore limb
bears scales to the base of the claws, the tail is massive and bears
no naked subterminal pad." In the former "there are 12 to 15
scales in the median dorsal row of the tail. . . . The scales of the
head, neck, shoulders, arm, and hind legs are dominantly dark olive-
brown. This color shades gradually into avellaneous over the dorsal
region. . . . Over the tail a deep Roman green assumes increasing
prominence in the apical part of the scale. . . . The species is hair-
less, except for a dense ring of short, circumorbital bristles and a
patch of similar hairs in front of the auditory meatus." The tail
182 EXTINCT AND VANISHING MAMMALS
length averages a little less than half of the total length. (Hatt,
19346, pp. 646-649.) The species attains a total length of 1,710 mm.
and a tail length of 830 mm. (Allen and Coolidge, 1930, p. 606) .
The range appears to extend from Sierra Leone and Liberia east
to the Ubangi-Shari Territory of French Equatorial Africa and
northeastern Belgian Congo. It corresponds rather closely to the
Upper and Lower Guinea Forest District of Chapin (1932, p. 90)
and of Bowen (1933, pp. 256, 258). "M. gigantea is known from the
West African Rain Forest and the adjoining wooded galleries" (Lang,
in Bequaert, 1922, p. 325). Matschie (1894a, p. 5) seems to extend
the range as far as Senegambia.
FIG. 20. Giant Pangolin or Scaly Anteater (Smutsia gigantea). After
photo by Lang.
Sierra Leone. A species of pangolin, said to be Smutsia gigantea,
is of fairly general distribution but is not commonly seen. It pro-
vides food for the natives. There is no evidence of depletion, and
no protective measures are taken. (Colonial Secretary's Office, in
litt., July, 1937.)
Liberia. This is a very rare animal in Liberia. A specimen
secured by Jackson at Cape Mount had consumed a large quantity
of termites and driver ants. The flesh is very tough and has a flavor
of formic acid. (Buttikofer, 1890, vol. 2, pp. 395-396.)
Another specimen is recorded by Jentink (1888, p. 56) from Little
Bassa.
A male of record size was obtained from natives at Paiata (Allen
and Coolidge, 1930, p. 606) .
Gold Coast. This species "is found in the savannah areas of
N. Ashanti and the Northern Territories.
"There is little doubt that all [the species of pangolins] are now
much less common than formerly, though it is probable that their
range has not decreased.
ORDER EDENTATA: EDENTATES 183
"Night hunting and the use of wire snares are the main causes of
depletion. Their meat is considered one of the greatest delicacies."
(Assistant Conservator of Forests, Gold Coast, in litt., July 22,
1937.)
Nigeria. The species is reported from Nigeria, but without a
definite locality record (Rosevear, 1937, p. 13) .
French Cameroons. It occurs in the forest region and is absolutely
protected except under scientific permit (Paris Agency, in litt.,
November, 1936).
Gabun. The Giant Pangolin is confined to the great forest. Only
a skin has come under personal observation. It was, however,
abundant at Mimongo in the region of Akelai. (A. R. Maclatchy,
in litt., February 5, 1937.)
Ubangi-Shari district, French Equatorial Africa. It appears to
be localized in the forested region. It is not threatened, and has
been totally protected since 1929. (L. Blancou, in litt., December,
1936.)
Belgian Congo. Schubotz (1912, p. 356) records a specimen from
Angu, on the Uele River.
Lang (in Bequaert, 1922, p. 320) says of the several local species
of pangolins:
"The signs of their fossorial practice are as often a cause of their
discovery as is the strong odor they emit, and dogs of native hunters
never fail to challenge their presence. Various highly valued talis-
mans, which their captors obtain from the claws, scales, hairs, and
other parts of some of the scaly ant-eaters, suffice to make them an
always welcome prize and their meat is an additional incentive for
their destruction."
Lang also describes (p. 325) a Pygmy method of capturing the
present species: "Pygmy boys, with one end of a strand of rattan
fastened to the waist and the other held by friends waiting outside,
entered the burrows without hesitation .... These boys, armed
only with a knife, merely fastened the rattan around the live pan-
golin, which they prodded from behind while their companions
pulled it out of the hole. These otherwise harmless beasts, when
touched while rolled up, suddenly switch their tail sidewise with such
force that, if one's hand is caught between the rough body scales
and the tail, it is seriously mutilated by the shearing action."
Lang records (p. 325) specimens from Bafuka, Niangara, Poko,
and Niapu in northeastern Belgian Congo.
"The only specimen I was ever able to obtain was dug out for
me by natives, with the expenditure of much labor and time, on the
Semliki side of the forest" (Christy, 1924, p. 228).
Schouteden states (1930, p. [95]) that the species ranges from
184 EXTINCT AND VANISHING MAMMALS
the Lower to the Upper Congo. He also records (1935, p. [62]) a
specimen from the Kivu region.
The several species of pangolins do not appear numerous any-
where in the Belgian Congo. The natives do not hunt them espe-
cially, and the Europeans not at all. Brush fires alone destroy a
great many. (A. J. Jobaert, in litt., November 10, 1936.)
South African Pangolin ; Scaly Anteater. I jzer Magauw ;
letermago (Boer)
SMUTSIA TEMMINCKII (Smuts)
Mani's temminckii Smuts, Enumeratio Mammalium Capensium, p. 54, pi. 3,
figs. 6-7, 1832. ("E regionibus, ultra Latakou sitis" = probably the region
north of Litakun, British Bechuanaland.)
FIGS.: A. Smith, 1849, pi. 7; Royal Nat. Hist., vol. 3, p. 229, fig., 1894-95;
Matschie, 1895, p. 143, fig.; W. L. Sclater, 1901, vol. 2, p. 217, fig. 148;
Fitzsimons, 1920, vol. 4, pi. facing p. 233.
Though widely distributed in South and East Africa, this seems
to be a decidedly scarce animal.
"General form somewhat elongated and lizard-like, covered every-
where, except on the lower surface of the head and body and inside
the limbs, with a series of over-lapping broad scales of a dark horn-
brown colour with paler edges and tips; head very small and
pointed," covered above with small scales; "no external ear . . . .
Across the middle of the back eleven rows of scales . . . ; limbs
short each with five toes and claws .... Tail very broad," cov-
ered above and below with 4-5 rows of scales. Head and body, 24
inches; tail, 18. (W. L. Sclater, 1901, vol. 2, pp. 217-218.)
"The scaly ant-eater is chiefly found to the north of the Orange
River, though said to occur rarely in Prieska and the other districts
just south of the river; from here it extends through the Orange
Free State, the Transvaal, Bechuanaland, the Kalahari and German
South-west Africa to Rhodesia; north of the Zambesi it occurs in
South Angola, Nyasaland and East Africa as far as Somaliland."
(W. L. Sclater, 1901, vol. 2, p. 218.) Matschie (1894, p. 5) extends
the range north to southern Kordofan (about lat. 17 N.).
"Well known in the Orange Free State (Ventersburg Albany
Museum) , the Transvaal, Bechuanaland, Ngamiland, and Southern
Rhodesia." Also "recorded from Northern Rhodesia, Nyasaland,
and according to Sclater Uganda, East Africa and Somaliland."
(Shortridge, 1934, vol. 2, p. 665.)
Cape Province and British Bechuanaland. At Litakun, British
Bechuanaland, Burchell (1824, vol. 2, pp. 423-424) observed a skin
lying on the hedge of a native cattle enclosure, "placed there . . .
to preserve the cattle from the evil effects of sorcery. . . . When-
ORDER EDENTATA: EDENTATES 185
ever a recent track is met with, the animal is traced to its hole and
dug out if possible, as the flesh, which is extremely fat, is esteemed
so great a delicacy that the law requires that every khaaka which
is killed shall be brought to the Chief."
Referring to this same general region, A. Smith says (1849, text to
pi. 7) : "Only one solitary specimen of this species was obtained by
the expedition before reaching 26 south latitude, and but two more
between that parallel and the tropic of Capricorn .... Its extreme
scarcity probably arises from its having long been zealously sought
after by the natives .... Whenever a specimen ... is secured,
it is immediately burned in some cattle pen, which, according to
the opinion of the sacrificer, tends to increase the health and fertility
of all cattle who may afterwards enter the fold. Not many years
ago a specimen was captured in the northern part of the Cape
colony."
"I have seen a dry skin from Upington " (Shortridge, 1934, vol. 2,
p. 665) . In Griqualand West the animal is now very scarce, and the
scales are used by the natives as medicine (McGregor Museum,
Kimberley, in litt., June, 1937) .
South-West Africa. In the Omaheke and the Kaukauveld it is
widespread but rather rare; skins are seen occasionally among the
natives. It is also reported by Bushmen in the Hukweveld. (Zu-
kowsky, 1924, p. 68.)
"The Pangolin occurs throughout South-West Africa," but is
"never plentiful. . . .
"It is apparently most numerous north of the Tropic of Capricorn
and in the sand-plains adjoining Bechuanaland.
"Rare in the vicinity of the Orange River and in the southern
parts of Great Namaqualand. . . .
"Pangolin scales (used as charms) were not infrequently seen in
the possession of Bushmen and other natives." (Shortridge, 1934,
vol. 2, p. 665.)
Angola. Monard (1935, p. 183) records two specimens from the
region between the upper Kului and the Kubango, where the natives
report the animal as rather common. Monard also mentions (p. 185)
specimens recorded by Bocage from Caconda and Mossamedes.
Transvaal. "A number of examples have been sent to the National
Zoological Gardens, chiefly from the Rustenburg and Marico districts
of the Transvaal" (Haagner, 1920, p. 237).
"ISIever very common and probably scarcer now as a result of
closer settlement. There is a demand for its scales by native witch
doctors for 'medicine/ as much as six pence per scale being paid,
so that this leads to a considerable amount of destruction of the
animal. Being entirely useful and harmless it should receive more
protection than is actually accorded it .... (Not included in the
186 EXTINCT AND VANISHING MAMMALS
game laws, i. e., without any special protection.) " (A. Roberts, in
Hit., November, 1936.)
Portuguese East Africa. Peters (1852, p. 174) records specimens
from Quitangonha, from near Cape Delgado, and from the vicinity
of Quelimane. He adds that the scales are made into finger rings
and worn as a protection against the "evil eye."
Kirk states (1865, p. 654) that it occurs near Sena.
Southern Rhodesia. Chubb (1909, p. 125) records a specimen
from AVankie, Matabelelend.
The species is by no means common. Most Rhodesian natives use
the skin as a charm, and for this reason the animals are in great
demand. Were it not for their retiring nature and strictly nocturnal
habits, they would be in danger of extermination. They will be
protected in the near future. (Game Warden, Wankie Game Re-
serve, in litt., March, 1937.)
Northern Rhodesia. Pitman notes (1934, p. 173) that this pango-
lin is "recorded from Batoka Province and Barotse." The natives
do not "seem to know of it in the areas I have toured (with the
exception of the Kafue Hook) ." He quotes Neave (1906) to the effect
that it is not unusual to see the scales worn as charms by natives
of the mid-Zambesi Valley.
Tanganyika Territory. Holmwood (1878, p. 632) records a speci-
men "from the coast opposite Zanzibar, lat. 6 S. ; but I have seen
what I took to be the same animal, both in Somali-land under the
equator and as far south as the Makna country opposite Mozam-
bique."
Matschie (1895, p. 143) records the animal from Wahumba,
Bagamoyo, Massai Nyika, and Mandera.
The Game Preservation Department (in litt., December, 1936)
reports no danger of extinction.
Kenya. The Game Warden (in litt., November, 1936) reports
no decrease, though the animal is not protected.
Three-cusped Pangolin; White-bellied Pangolin; Pale-bellied
Pangolin; Pointed-scaled Pangolin. Pangolin tricuspide
(Fr.). Dreizackige Schuppentier (Ger.)
PHATAGINUS TRICUSPIS (Rafinesque)
Manis tricuspis Rafinesque, Annales Gen. Sci. Physiques [Bruxelles], vol. 7,
p. 215, 1821. (Type locality not stated; restricted by Allen and Coolidge
(1930, vol. 2, p. 606) to "West Africa.")
FIGS.: Royal Nat. Hist., vol. 3, p. 230, fig., 1894-95; Johnston, 1906, vol. 2,
p. 749, fig. 292; Schouteden, 1930, p. [88], fig. 1; Hatt, 19346, pis. 36-37,
and 1934c, p. 727, lower fig., p. 730, upper fig., p. 731, right-hand fig.;
Rosevear, 1937, p. 12, fig. 1.
ORDER EDENTATA: EDENTATES 187
This species occurs in apparently larger numbers than the other
African pangolins.
It is "an arboreal species with a tail constituting over half the
total length. The characters of the tail tip and the fore limbs are
like those of Manis longicaudatus. The scales, however, are small
and numerous, brown, and during mid-life, tricuspid. The post-
scapulars are not enlarged." Under parts grayish white. In young
animals "the margins of the scales are even, but with ensuing wear
. . . the scales become sharply dentate, or, usually later, tridentate."
In old age the animals have "cuspless, worn, elongate scales." In
half-grown and mature animals the unsealed parts of the skin are
covered with hair, attaining a length of 20 mm. Longitudinal rows
of scales, 21-25; marginal caudal scales, 35-40. Total length, up to
1,027 mm.; tail, 607 mm. (Hatt, 19346, pp. 655-658.)
This pangolin is not confined to the Upper and Lower Guinea
Forest Districts but ranges southward into the Southern Congo
Savanna District and eastward into the Uganda-Unyoro Savanna
District of Chapin (1932, p. 90) and of Bowen (1933, pp. 256, 258).
Hatt (1934b, p. 656) records specimens from Liberia, the Ivory
Coast, Cameroons, Fernando Po, Gabun, the lower Congo, Kasai
district, and central Angola. According to Matschie (1894a, p. 6),
the range extends west to Gambia, and Jentink (1882, p. 208) has
a record from Sierra Leone.
Liberia. The species appears to be distributed over the entire
region. Specimens are recorded from Buluma, Schieffelinsville,
Junk River, Hill-town, and Farmington River. The animal can be
tamed and kept a long time in houses, where it runs free and preys
upon ants, cockroaches, and other troublesome insects. (Buttikofer,
in Jentink, 1888, p. 57.)
Allen and Coolidge (1930, vol. 2, p. 606) record "a native-made
skin bought at Since."
Gold Coast. Hayman (1936, p. 937) records specimens from
Goaso and Mampong.
The species is found through much of the forest country, but is
doubtless much less common now than formerly (Assistant Con-
servator of Forests, Gold Coast, in litt., July 22, 1937) .
Fernando Po. Fraser (1848, text to pi. 28) records the species
from this island, where "the flesh is said to be exceeding good eating,
and is in great request among the natives."
Gabun. This pangolin is confined to the great forest. Although
legally protected, it is actively hunted by the forest natives, who
capture great quantities. To prevent this is difficult, for the animal
is taken in trigger traps set for small game. The real safeguard
would be the prohibition of this type of trap; but those who know
188 EXTINCT AND VANISHING MAMMALS
the brush know how much such prohibition would be worth. (A. R.
Maclatchy, in litt., February 5, 1937.)
French Equatorial Africa. Matschie (1894a, p. 6) records the
species from Loango.
It is common almost everywhere in the Ubangi-Shari district.
It has been totally protected since 1929, and is not threatened. (L.
Blancou, in litt., December, 1936.)
Angola. Monard (1935, p. 185) quotes Bocage to the effect that
this pangolin is rather common at Bembe and Malange; he also
gives records for Bimbi and Cazengo.
Belgian Congo. Schwarz (19206, p. 1061) records specimens
from Libenge on the Ubangi, Panga on the Aruwimi, Angu on
the Uele, and Avakubi on the Ituri; also from Kudurma and Ka-
bayendi in the Niam-Niam country (not far from the Congo-Sudan
boundary) .
Lang (in Bequaert, 1922, pp. 320-323) remarks that tricuspis
is the commonest of the African pangolins. "Being timid, they
readily make use of their natural safeguard and, when even slightly
annoyed, roll up in a ball .... When forcibly unrolled, they
may succeed in driving off their tormentors by well directed jets of
an ill-smelling, acrid liquid from the anal region; native dogs suffer
for a considerable time from the effect of this substance, which
greatly irritates their mucous membranes. . . .
"If unmolested and placed near their favored prey, they uncoil
readily .... One soon realizes how thoroughly they are special-
ized as ant-eaters, for their methods of attack and disposal of ants
are as effective as their ways of guarding themselves against the
defensive means of their prey. In the regions we visited, the pan-
golins preferred true ants, as stomach contents clearly showed,
though many of our captives would plunder termitaria with great
eagerness. . . .
"One taken near a column of army ants (Dorylus) merely made
good its escape, another quickly broke up the well-ordered line.
. . . Lashing its sticky tongue through the confused crowds, the
ant-eater lost no time in moving back and forth along the ant
column as quickly as the dense clusters vanished into its mouth.
Its hunger satisfied, it at once retreated, freeing itself of the few
army ants that had managed to dig their mandibles into the soft
parts of its hide. M. tricuspis fed freely on many other kinds of
ants. Those we had alive at Avakubi, Medje, and Niapu were
particularly fond of ants of the genus Myrmicaria. . . .
"African pangolins have helped to enrich the stores of witchcraft."
Hatt (19346, p. 645) records 66 specimens from Akenge, Avakubi,
Faradje, Gamangui, Medje, Ngayu, Niangara, Niapu, Poko, and
Stanleyville.
ORDER EDENTATA: EDENTATES 189
" Uganda. An arboreal pangolin (presumably tricuspis) is reported
by Johnston (1902, vol. 1, pp. 395-396).
"Two or three species occur in Uganda, the common representa-
tive being Phataginus tricuspis, a forest species. There is no reason
to believe that Pangolins are any less plentiful than formerly. In the
Mabira Forest P. tricuspis is abundant. All species of Pangolins
are completely protected in Uganda." (Game Warden, Uganda,
in litt., December, 1936.)
Long-tailed Pangolin; Black-bellied Pangolin. Pangolin a
longue queue (Fr.). Langschwanzige
Schuppentier (Ger.)
UROMANIS LONGICAUDATA (Brisson)
Pholidotus longicaudatus Brisson, Regne animal, vol. 3, Quadr., p. 19, 1762.
("Probably West Africa" (Allen and Coolidge, 1930, p. 606).)
SYNONYMS: Manis teiradactyla Linnaeus (1766); Manis macroura Erxleben
(1777); Manis hessi Noack (1889).
FIGS.: Noack, 1889a, pi. 1; Johnston, 1906, vol. 2, p. 753, fig. 295; Bequaert,
1922, pi. 25, fig. 1; Allen and Coolidge, 1930, pp. 603-605, figs. 447-449;
Schouteden, 1930, p. [94], fig. 3a; Halt, 19346, pi. 35, figs. 1-2, and 1934c,
pp. 726 (both figs.) and 731 (lower fig.) ; Rosevear, 1937, p. 12, fig. 3.
The very limited amount of information available concerning
this species suggests that it is one of the rarest of the African
pangolins.
This is "an arboreal species with a long prehensile tail, equaling
about two-thirds of the total length. . . . The forearms bear no
scales, but are covered with hair. The scales are large, yellow, and
on the flanks are keeled. The two inferior postscapular scales are
markedly larger than those adjacent to them. . . . The belly hair
is black in most individuals .... The whole face . . . dark brown,
nearly black." Total length up to 937 mm.; tail, 645. (Hatt, 19346,
pp. 651-652.) Thirteen rows of scales on the body; 44 marginal
scales on the tail; two rows of 9-10 scales before the tail tip (Mat-
schie, 1894a, p. 7).
The range appears to be more or less coextensive with the Upper
and Lower Guinea Forest Districts of Chapin (1932, p. 90) and of
Bowen (1933, pp. 256, 258). Jentink (1882, p. 207) records speci-
mens from as far west as Senegal and Sierra Leone. Otherwise the
species is known from Liberia to Gabun and the northeastern Bel-
gian Congo.
Liberia. The animal is pretty rare, though a number of living
specimens were received, including one at Soforeh Place. (Butti-
kofer, 1890, vol. 2, pp. 393-394.)
Jentink (1888, p. 56) records additional specimens from Hill-town
and Farmington River.
190 EXTINCT AND VANISHING MAMMALS
Live specimens were brought to Allen and Coolidge (1930, vol. 2,
p. 606) at Lenga Town on the Farmington River and at Paiata.
Gold Coast. Specimens are recorded from Dabocrom and Elmina
(Jentink, 1882, p. 207) ; also from Goaso (Hayman, 1936, p. 937).
The species is found through much of the forest country, but there
is little doubt that it is now much less common than formerly
(Assistant Conservator of Forests, Gold Coast, in litt., July 22,
1937).
Cameroons. Hatt (19346, p. 653) records the species from this
country, without stating the exact locality.
Gabun. Hatt (19346, p. 652) records a specimen from Fernand
Vaz.
Belgian Congo. Noack (1889a, p. 100) based his name Manis
hessi upon a specimen from the vicinity of Banana, at the mouth
of the Congo.
Hatt (1934b, pp. 651, 653) records specimens from Bolobo and
Lukolela on the Lower Congo, and from Akenge, Gamangui, Medje,
and Niapu in the northeastern part of the country.
Order RODENTIA: Rodents
Family LEPORIDAE: Hares and Rabbits
This family is of nearly cosmopolitan distribution; but it is
absent from Madagascar and part of the Malay Archipelago, and it
was lacking in Australia until introduced. There are about 11 genera
and over 200 species and subspecies. There is generally an abun-
dance of individuals, and only a single species, the insular Amami
Hare, comes within the scope of this report.
Amami Hare; Liu Kiu Hare
PENTALAGUS FURNESSI (Stone)
Caprolagus furnessi Stone, Proc. Acad. Nat. Sci. Philadelphia 1900, p. 460,
1900. ("Liu Kiu Islands.")
This remarkable insular hare, unique representative of its genus,
has been suitably recognized and protected by the Japanese Govern-
ment as a "Natural Monument."
Size approximately that of Lepus americanus; hind foot, tail,
and ears remarkably short ; claws very large and strong ; soft under-
fur plumbeous ; long hairs coarse and hispid, brownish black, many
with buff annulations; a median black stripe from neck to rump;
under parts mostly pale buff. Total length of flat skin, 550 mm.;
tail, about 8 mm., (Stone, 1900, pp. 460-461.) "Pentalagus is the
most marked of any of the genera of the Leporidae, the tooth
ORDER RODENTIA: RODENTS 191
formula, the structure of the teeth, the relative size of the radius and
ulna, and the very short tarsus and metatarsus being peculiar to
the genus and unlike anything in the rest of the family" (Lyon, 1904,
p. 430).
In the original description Stone (1900, p. 460) records two speci-
mens.
Thomas (1906a, p. 357) records a specimen from "Oshima, Oki-
nawa, Liu-Kiu Is.," and adds: "Another specimen is now living in
the Duke of Bedford's menagerie at Woburn."
"The distribution of this species ... is restricted to the Islands
of Amami-Oshima and Tokuno-shima in the Loochoo archipelago
where it is endemic" (Kaburaki, 1934, p. 4183) .
"Number is unknown, but as it is carefully protected as one of the
'Natural Monuments/ by the Law for Preserving Scenery, Historic
and Natural Monuments, and it is also strictly prohibited to capture
the species without special permission, and besides it is forbidden
by the game law, it will never become extinct" (Nagamichi Kuroda,
in litt., July 5, 1938) .
Family CASTORIDAE: Beavers
The single genus of this family is repres'ented by one species
(Castor canadensis) , with 20 subspecies, in North America, and by
another species (fiber) , with perhaps four subspecies, in Europe and
northern Asia. All the American forms have been treated by Dr.
Allen in the preceding volume (1942), and an account of Castor
fiber and its subspecies follows here. It is primarily the demands
of the fur trade that have brought about trie deterioration in the
status of the Beavers.
European Beaver. Castor; Bievre (Fr.) Biber (Ger.)
CASTOR FIBER Linnaeus
[Castor] fiber Linnaeus, Syst. Nat., ed. 10, vol. 1, p. 58, 1758. (Sweden.)
FIGS.: Geoffrey and Cuvier, Hist. Nat. Mamm., vol. 6, pi. '275, 1824; Brandt
and Ratzeburg, 1829, pi. 3; Blasius, 1857, p. 403, fig. 224; Royal Nat. Hist.,
vol. 3, p. 97, pi., 1894-95; Collett, 1898, pi. 12; Martin, 1910, pi. 10;
Didier and Rode, 1935, p. 188, fig. 98.
While treated here as a specific unit, Castor fiber has been divided
into a number of forms, including the following which are con-
sidered by Kuntze (1935, p. 64) to be more or less tenable:
C. /. fiber Linnaeus (Sweden) ;
C. f. vistulanus Matschie (western Poland) ;
C. /. albicus Matschie (Dessau, Anhalt, Germany) ; and
C. /. galliae Geoffrey (the Rhone, France).
192 EXTINCT AND VANISHING MAMMALS
The former range of the Old World Beaver included the forested
regions of Europe and northern Asia. The original colonies are now
extinct in all save a few localities in France, Germany, Norway,
Poland, Russia, and Siberia. In recent times the animal has been
reintroduced into England, Sweden, and Latvia.
The general form is heavy and thickset; eyes and ears small;
hind feet large, broad, and webbed, the claw of the fourth digit
with a horny, compressed supplement; tail scaly, mostly naked,
broad, depressed ; general color a peculiar and very uniform clayey
buff, the under parts a little more yellowish. Head and body (fe-
male) , 820 mm. ; tail, 380 mm. (Miller, 1912, pp. 948-952.) Weight,
15 to 25 or even 36 kilograms (Didier and Rode, 1935, p. 188).
Great Britain. In Wales, in A. D. 940, Beaver hides were req-
uisitioned for making the borders of the king's garments; it was
evidently then a rare animal. In 1188 it was still found on at least
one river in Wales and on a single river in Scotland, though it had
apparently died out quite generally in other parts of Great Britain.
"The written records we have of its occurrence are very frag-
mentary, and not wholly satisfactory." Remains have been exhumed
in both England and Scotland. Various place names in England
indicate the former occurrence of Beavers there. (Harting, 1880,
pp. 33-46.)
Their skins were exported from England and Scotland until the
middle of the twelfth century. Beavers x were reintroduced on the
island of Bute, Scotland, in 1874, but died out about 1890. There
were similar introductions in Suffolk, England, in 1870, and in Sussex
at some time prior to 1905. (Millais, 1905, pp. 162-163.)
In 1663 a good Beaver hat in England cost 85 RM. in German
currency (Kriiger, 1931, p. 54).
Spain. Strabo, writing of this country in the first century B. C.,
is said to mention the Beaver as a well-known animal (Blasius, 1857,
p. 407; Kriiger, 1931, p. 52).
France. The Beaver was once widely distributed in France,
being found on many watercourses in various basins. It gradually
became rare, but in the sixteenth century was still found on many
rivers, principally the Oise, the Somme, and the Marne. Today it
is found only on the Rhone and its tributaries, below Valence. The
principal habitats are: (1) the mouth of the Ardeche; (2) the
mouths of certain small watercourses the Ceze, the Tave, and the
Aigues; (3) the vicinity of Roquemaure and the lie de Miemas;
(4) the vicinity of Avignon and the lie de Barthelane; (5) along the
course of the Garden; (6) between Tarascon and Beaucaire; (7) on
i According to Kriiger (1931, p. 53), Scottish importations at this period were
of Canadian Beavers.
ORDER RODENTIA: RODENTS 193
most of the course of the Petit-Rhone, in rather numerous colonies ;
(8) on the Grand Rhone, beginning at Aries, in less numerous
colonies.
The 60 or 70 known stations are certainly not the only ones. A
rough estimate of the total population is 300 individuals.
The reasons for depletion are numerous and diverse. Although
the Beaver was always hunted for its valuable fur, it was long
considered, up to recent years, as harmful and thus was under official
ban. Trapping in submerged nets has been particularly fatal.
The frequent floods on the Rhone have been a serious factor in
destruction.
FIG. 21. European Beaver (Castor fiber). After Brehm.
Perhaps pollution of the watercourses in certain areas is to be
blamed, for autopsies have revealed tubercular lesions. It does not
appear from the autopsies, however, that the fecundity of the species
has been diminished.
When its existence was threatened a dozen years ago, the warning
issued by certain naturalists rapidly bore fruit. Restrictions on hunt-
ing, establishment of reserves, warden service, propaganda in favor
of the Beaver, and appreciation of this rare animal by the local
population, have been effective in its conservation. Its. future seems
brighter, and in general the colonies seem more prosperous than a
little while ago. It would be easy to improve the situation by the
creation of more reserves, by the establishment of zones of refuge
at the time of floods, and by the repopulation, if possible, of old
abandoned colonies. (E. Bourdelle, in litt., March 6, 1937.)
Since 1909 the hunting and capture of Beavers have been pro-
hibited for all time. Twenty kilometers of the Rhone have been
194 EXTINCT AND VANISHING MAMMALS
declared a protected reserve. As a result of persecution the species
has lost the habit of constructing dams and lives in burrows in the
banks of the streams. (Didier and Rode, 1935, pp. 192-193.)
Trouessart wrote in 1884 (pp. 119-121) that its northern limit
in the Rhone Basin was approximately Valence (as it is now). He
added that it was becoming rarer each year, owing to relentless per-
secution. Also that its flesh is excellent, and since it is hunted for
its hide and for its castor, as well as because of its depredations in
young plantations, its early exterminaton in France could be pre-
dicted. But such a fate has been happily warded off.
A few years ago its castor was worth more than 250 francs a
pound. For a long time the Syndicat des digues du Rhone paid a
bounty of 15 francs on Beavers, on account of the alleged damage
to dikes. But with better information the bounty was abandoned.
(Martin, 1910, p. lOb.)
Italy. Gesner (1551, vol. 1, p. 337) mentions the Beaver as
occurring at the mouth of the Po.
Yugoslavia. The species is entirely exterminated, the last speci-
mens having been observed in 1859 at Syrmia on the Danube and
at about the same time in Bosnia on the River Ukrina. A good
many fossil remains have been found in Croatia and Slovenia. (M.
Hirtz, in Hit., December, 1936.)
Rumania. The species existed in Transylvania up to about 1500,
and in Moldavia to 1823 (Calinescu, 1931, p. 82).
Hungary. Extermination took place in the first half of the nine-
teenth century (J. Schenk, in litt., November, 1936).
Austria. In 1867 Beavers still lived northwest of Salzburg, where
the Sur discharges into the Salzach, but by 1870 only disused
burrows could be found. There was formerly a protected colony
in the plains of the Danube near Wien. But now the Beaver is no
more to be found in the Danube region. In 1861 the castor fetched
600 Gulden in Salzburg. (Kruger, 1931, pp. 52-54.)
Czechoslovakia. Under the protection of the Princes of Schwarz-
enberg, Beavers survived long in the tributaries of the Moldau, but
the last of them died in 1883 (Kruger, 1931, p. 53).
Switzerland. Gesner (1551, vol. 1, p. 337) reports the Beaver
as a very common animal in the large rivers. But it could not sur-
vive strong persecution (Kruger, 1931, p. 52). Millais (1905, p. 160)
quotes Harting to the effect that "Beavers were to be found in the
Aar, the Linnet, and the Reuss, and up to the last century [eight-
eenth] a few still lingered on the banks of the last-named stream,
on the Thiele, and the Byrse."
Germany. On the Rhine the animals died out more than 300
years ago. In Westphalia they occurred up to the middle of the
nineteenth century, and the very last was killed apparently in 1877.
ORDER RODENTIA: RODENTS 195
In the Province of Hannover they disappeared more than a hundred
years ago.
Beavers now find refuge only on the middle course of the Elbe
between Wittenberg and Magdeburg, together with its tributaries
the Mulde, Saale, and Nuthe, and the adjacent Altwasser. The
population was estimated in 1890 at 200; in 1913, at 188; in 1919, at
42; in 1922, at 200; in 1926, at 164; in 1929, at 263.
The almost total destruction of the Beaver in Europe is to be
explained only by avaricious persecution. It was pursued because of
its tasty flesh, its valuable pelt, and especially its castor, which
commanded a very high price as a panacea.
The presence of the Beaver today on the middle Elbe is due to
certain protective measures. Formerly it enjoyed no protection.
The Prussian Game Law of 1907 gave it a 10-month closed season.
In 1921 and again in 1929 it was given complete protection. Along
the Elbe mounds are constructed to furnish a refuge during floods,
and some willow plantations are provided as food. There are re-
strictions on fishing and trapping in the immediate vicinity of the
Beaver burrows. The Provincial Assembly has made an appro-
priation of 1,000 RM. for settlement of claims for damage by
Beavers. Despite these protective measures, the Beaver stand in-
creases only slightly or not at all.
Owing to the penalties involved and the difficulty of disposing
of the skin, deliberate killing for profit has practically ceased.
But some animals still fall victims each year to the human lust for
killing. More serious is the killing for protection against damage.
In the eyes of the country people and the fishermen the Beaver is
injurious. It steals their potatoes and turnips and destroys their
fruit and forest trees. Some are caught and drowned in fish nets
and traps. They are also endangered by steel traps set for Otters.
Floods and drifting ice constitute the greatest menace to the
Beaver. Tuberculosis was found in a dead animal.
Introduction of Beavers into other parts of Germany, where they
may find suitable living conditions and safety, is being considered.
(Kriiger, 1931, pp. 53-56.)
The present range in Germany is on the Elbe between Torgau
and Magdeburg, and on the adjacent tributaries. Tuberculosis has
been found the cause of death of a number of animals. Some Beavers
have been introduced in the Schorfheide, near Berlin. (Reichsstelle
fur Naturschutz, in litt., October, 1936.)
In Wiirttemberg the last specimen was killed in 1869 on the
Danube at the mouth of the Iller River (Wiirttembergische Natur-
aliensammlung, in litt., October, 1936) .
Blasius refers (1857, p. 407) to the former occurrence of Beavers
in northwestern Germany on the Moselle, the Maas [now in the
196 EXTINCT AND VANISHING MAMMALS
Netherlands], and the Weser; in the Luneburg area; and on the
Schunter near Braunschweig. He also speaks of their more recent
occurrence on the Havel and the Oder, in the Altmark, on the
Vistula [now in Poland], in East Prussia, and at Schwenckfeld in
Silesia.
"Harting says that 'at the close of the last century [eighteenth]
many localities are reported to have been frequented by Beavers,'
notably in Altmark, Preignitz, Middlemark, on the rivers Spree and
Haxel and in the vicinities of Berlin, Potsdam, Oranienburg, Lieben-
walde, Trebbin, Nauen, and Konigshorst. . . . Wagner in 1846
mentions Beavers as living on the Danube, Amper, Isar, Iller,
Salzach, and the Oder." (Millais, 1905, p. 161.)
Denmark. The species was formerly distributed all over Den-
mark, including Bornholm (Winge, 1908, p. 96) .
Norway. Trade in Beaver skins was carried on early in the
Middle Ages. Probably most of the Norwegian furs were exported
to England. The species had begun to decrease by the close of the
seventeenth century. In the middle of the eighteenth century it was
probably still distributed throughout most of the woodland valleys,
from the southernmost parts of the country to the farthest confines
of Finmarken.
In 1896 its range was chiefly in the Stifts of Christiania and
Christiansand. The largest colony was located on the Nisser River
in Nedenaes Amt.
The trees felled are used both for food and for building material.
The Beaver prefers the aspen (Populus tremula) and after that the
birch, oak, and alder.
According to an old superstition, the castor has the power to
frighten away whales approaching a boat. In some parts the castor
is worn from the garter as a specific for worms. In the sixteenth
century the tail was regarded as a table delicacy. The teeth are
worn as amulets in Finmarken, partly for ornament, partly as a
protection against sickness, and were offered to the gods at the
place of sacrifice and buried in the graves of heathen Lapps.
By the middle of the nineteenth century the Beaver was fast
becoming extinct, but the Game Laws of 1845 checked the decrease.
By the end of the century it was on the increase and had extended
its range by migration. The number was estimated at about 60 in
1880, about 100 in 1883, and perhaps a few more in 1896. By this
time Norwegian Beaver skins were very rarely in the market, and the
castor was of little value.
For a period of about 40 years after 1855 Beavers were allowed to
be killed under certain restrictions. Then a closed season of 10 years
was declared for the entire Amt of S0ndre Bergenhus and for the
whole of Aamli. (Collett, 1898, pp. 105-126.)
ORDER RODENTIA: RODENTS 197
By 1931, as a result of state protection, the numbers on the Nidelf
River had increased to 12,000 (Kriiger, 1931, p. 53).
A limited amount of hunting is now allowed. During late years
Beavers have been transplanted to the northern parts of the country,
where they seem to thrive well. In some districts the farmers com-
plain of damage to the forests by the Beavers. (Hj. Broch, in litt.,
December, 1936.)
Sweden. The Beaver was formerly distributed all over the
country, but gradually it became extinct. The last specimen in
Smolandia was probably killed about 1800; that in Jemtland prob-
ably about 1870. It was the value of the fur and the castor that
caused its extermination. Beavers of the same race from Norway
were first introduced in Jemtland in 1922, in Westerbotten in 1924,
and in Wermland in 1925. These have all increased, and the number
in Jemtland is now estimated at several hundred. There have also
been introductions in other provinces, and further trials of this kind
are planned. The animal is now very popular in Sweden, and its
future appears to be rather promising, especially since it is protected
throughout the year. (Einar Lonnberg, in litt., 1937.)
Latvia. In former times the Beaver was found on many of the
smaller rivers, but owing to excessive hunting it was exterminated
about 1870. In 1927 the Government introduced four Norwegian
Beavers in the State Forest of Kurland, and in 1936 two others in
Smiltene in Livland. They have now increased in number to about
40. Hunting is forbidden. (Forest Department of Latvia, in litt.,
March, 1937.)
Lithuania. The species is probably exterminated. Since the
World War two specimens have been illegally captured: one in 1921
on the Dubisa, and one in 1935 on the Nemunas. Hunting is for-
bidden. Reintroduction from neighboring countries is desirable. (T.
Ivanauskas, in litt., November, 1936.)
Poland. Game protective measures were instituted as early as
the beginning of the eleventh century, when Boleslaus I the Great
proclaimed an act for the protection of the Beaver (Benedyct Ful-
inski, MS., 1933) .
In ancient times and perhaps even in the seventeenth century
the Beaver was quite common in all Poland. Owing to the reduction
of forest areas and especially to the regulation of rivers, it retired
to the eastern and northeastern parts of Poland, where it is still
found. (M. Siedlecki, in litt., October, 1936.)
Three preserves in the state forests, aggregating 684 hectares, are
sanctuaries for Beavers. Another, the Bucharzewo Preserve, of about
5 hectares, contains Canadian Beavers. (Benedyct Fulinski, MS.,
1933.)
198 EXTINCT AND VANISHING MAMMALS
The present colonies of Castor fiber vistulanus all lie in the river
basins of the Niemen, the Pripet, and the Dnieper, and perhaps also
the western Duna. A map shows the present distribution at 14
stations in the Niemen Basin and at 16 stations in the Pripet Basin.
The last records on the middle course of the Vistula were in 1850;
on the headwaters of the Vistula system, in 1861 ; on the headwaters
of the Bug, in 1861 ; on the headwaters of the Dniester, in 1851. The
largest colony in Volhynia in 1928 was estimated to contain 100
individuals. In 1928-29 the total number in Poland was estimated
at 235. The animals are now very strictly protected. (Kuntze, 1935,
pp. 65-68.)
Russia. Various early records are summarized by Nehring (1890,
p. 105) as follows: Pallas (ca. 1770) reported that Cossack hunters
sought Beavers on the steppe rivers of the Samara region, where the
animals occurred very sparingly. According to Rytshkov, Beavers
still existed in 1760-70 in the Bashkiri region; according to Evers-
mann, about 1850 in the Perm Government; according to Kessler,
at the same period in many rivers of the Kiev and Poltava Govern-
ments; according to Krynitzki, about 1835 near Kherson on the
lower Dnieper.
Trouessart (1910, p. 130) includes northern Russia and southern
Russia (Caucasus and rivers of the Caspian) in the range of the
species. Millais states (1905, p. 161) that "Beavers were found on
the Petchora and the Dwina in Russia until 1842, and possibly a few
may still exist in their unfrequented tributary stream-."
Of Russian Beavers we know comparatively little. In 1884, 566
individuals were counted in the Rokitno Swamps. But by the time
of World War I this number had greatly decreased, despite protec-
tive measures instituted in 1911. It is doubtful if the colony set
out on the Voronesh in 1886 still exists. (Kruger, 1931, p. 53.)
The species was formerly widely distributed in the forested areas
but is now almost exterminated and exists only in some reserves.
These are in the Ukraine (on the Rivers Teterev, Soge, and Desna,
on the tributaries of the Pripet, and in the former Government of
Chernigov) ; in the Western Area and in White Russia ; and on the
Usman in the former Government of Voronesh. In 1935 the total
number of Beavers in the U. S. S. R. (including Siberia) was esti-
mated at 2,500-3,000. (W. G. Heptner, in Hit., December, 1936.)
Siberia. According to Eversmann (as reported by Nehring, 1890,
p. 105), Beavers still existed about 1850 in the Baraba Steppe
(between the Irtish and the Ob Rivers). "Gone from the Yenisei
and Irtish, where formerly they were common, they were reported
from the Pelyn, a tributary of the Obi, in Western Siberia, until
1876, and they may still exist there" (Millais, 1905, p. 161). Troues-
sart (1910, p. 130) includes Turkestan as well as Siberia in the
ORDER RODENTIA: RODENTS 199
range of the species. According to Millais (1905, p. 161), it was
found even "as far east as Behring Straits." Kriiger states (1931,
pp. 52-53) that it once ranged from the Urals to the Pacific Coast,
but that the white examples with yellowish backs on the farther
side of the Urals probably have not survived, and that there are
reports of the complete extirpation of the species in Siberia. Ac-
cording, however, to W. G. Heptner (in litt., December, 1936), it
exists on the Rivers Konda and Sosva and their tributaries in the
Ob Basin.
""Schrenck (1859) reported the Beaver from Sakhalin, and he is
quoted as authority by Aoki (1913, p. 298) and by Hatta (1928,
p. 1036). The record is questioned, however, by Kuroda (1928,
p. 224) , who calls attention to the lack of specimens.
The paucity of beaver records from the Siberian wilderness sug-
gests that the animal may never have been very abundant or thor-
oughly distributed over that country. Perhaps the Siberian taiga,
with its predominant coniferous growth, does not provide a sufficient
quantity of the Beaver's favorite food trees, such as the aspen and
other deciduous species, to support the animal in large numbers.
Turkey, Syria, and Iraq. Danford and Alston (1880, p. 60) give
an unsubstantiated report of a beaverlike animal in the marshes
between Kaisariyeh and Indjesu, Turkey. They also cite reports by
Heifer and Heifer (1879) of Beavers on the Euphrates near Aleppo,
and by Schmarda (1853) in Mesopotamia.
Persia. "The beaver, according to Eichwald, is common in the
Araxes .... I insert it in the Persian fauna with some doubts."
(Blanford, 1876, p. 51.) The above-mentioned report from the
Araxes is categorically denied by later authors (Satunin, 1906,
p. 374).
Mongolia. In the upper Yenisei Basin, Tannu-Tuva, "a few
beavers still exist in the upper tributaries of the Bei-Kem; but
they are very rare, and their skins are seldom brought down to
the markets. In old days they were mentioned as being included
in a tribute sent by the Khan of the Ubsa region, then paramount
chief of the Uriankhai tribes, to the Czar of Russia." (Carruthers,
1913, p. 228.)
The species "still exists . . . , it is said, in the highest tribu-
taries of the Black Irtish in the Mongolian Altai" (Carruthers, 1913,
pp. 630-631).
Manchuria. "An animal recorded by Schrenck, but which does
not appear to belong to the Manchurian fauna, is the beaver (Castor
fiber) . It is true that skins of this animal have been secured from
the natives in the Amur region, and that they find their way to the
fur market in such places as Harbin and Mukden in Manchuria, but
recent investigation tends to show that these skins have been brought
200 EXTINCT AND VANISHING MAMMALS
from Alaska, having been bartered from one tribe of natives to
another till they came into the hands of Russian or Chinese fur
traders." (Sowerby, 1923, p. 170.)
India. The Beaver may have occurred even in India, since, ac-
cording to Buffon, the religion of the Magi forbade them to kill
this animal (Blasius, 1857, p. 407) .
Egypt. Since the species is supposed to be represented in the
Egyptian hieroglyphs, it may have occurred in Africa (Blasius,
1857, p. 407).
Economics. "Had not the use of its hair in the manufacture of
hats been superseded by that of silk, there is little doubt that the
beaver, both in the Old World ^nd in America, would by this time
have been numbered among extinct animals. As it is, the creature
has but a hard time of it at best, for although there is no longer a
demand for its hair by the hat-manufacturer, yet beaver-fur is an
article highly valued by the furrier, and equally highly esteemed
by the fair sex." (Lydekker, 1903, p. 244.)
Family CRICETIDAE: Hamsterlike Rodents
While various authors are not in accord on the limits of this family,
it is probably safe to say that it consists of more than a hundred
genera and more than a thousand forms. Representatives occur over
the greater part of the world, and in general their numbers are legion.
However, 14 New World forms are included in the preceding volume
by Dr. Allen (1942) , and the 6 forms of the African genus Lophiomys
are discussed here. A recent authority (Ellerman, 1941) makes a
separate family (Lophiomyidae) of this genus.
Genus Lophiomys Milne-Edwards: African Maned Rats
The following remarks of the Committee of Experts (Hemming
et al, 1938, p. 13), while naming only a single species, may be
taken to apply to all known forms of this rare and peculiar genus:
No species of rodent was included in either class of the Annex in the
Convention of 1933, presumably owing to the small size and insignificant
appearance of the majority of the species involved, and to the fact that they
do not fall into the category of game animals. We see no reason however
why a species of this Order should not be placed in the Annex if owing to
their rarity they are in danger of extinction.
A species of this Order which we should like to see protected is the Crested
Bush Rat, Lophiomys imhausi Milne-Edwards, a very remarkable species
living at altitudes of between seven and nine thousand feet in the mountains
of Abyssinia and Kenya. This species which lives in pairs in dead trees and
similar cover is peculiarly liable to attack and its numbers are known to have
diminished considerably in recent years.
We accordingly recommend that this species should be included in Class A
of the Annex which, owing to the fact that rodents are not game animals,
ORDER RODENTIA: RODENTS 201
appears to us the most appropriate method of securing their protection. In
the case of this particular species we shall hope to have received before the
next meeting of the Conference the data to be collected by the Italian Scien-
tific Mission.
Hollister remarks (1919, p. 37) : "Although a few specimens of
the maned rat find their way into collections from time to time,
the animal is still so rare that no suitable series are available for
study. If all the collections in various museums were combined it
t would still be impossible to form any correct idea of the relation -
'ships of the named forms, and it will doubtless be many years before
sufficient material has accumulated."
Under these circumstances the classification and nomenclature
in the following accounts of the known forms of Lophiomys must be
considered as no more than provisional. Possibly all the forms so
far described will eventually prove to be no more than subspecifically
distinct. All exhibit the same general color pattern.
Sudan Maned Rat
LOPHIOMYS AETHIOPICUS (Peters)
Phractomys aethiopicus Peters, Zeitschrift Gesammten Naturwissens. Halle,
vol. 29, p. 195, 1867. (Based upon a skull from Maman, north of Kassala,
Anglo-Egyptian Sudan.)
FIG.: Anderson and de Winton, 1902, pi. 51.
This species seems to be known chiefly from single specimens
collected at no more than about half a dozen localities in the Anglo-
Egyptian Sudan and Eritrea.
Under the name of L. imhausi, Anderson .(in Anderson and de
Winton, 1902, p. 289) describes an adult female from Erkoweet(?),
on the mountains between Suakin and Sinkat, Sudan, somewhat as
follows: Denser fur generally gray at the base, with a broad white
band and wide brown tip; the long hairs broadly tipped with white;
a triangular white area on top of the head, prolonged backward
below the ears to the side of the neck, where the adpressed lateral
band of yellowish hairs commences; a white spot below the eye;
front and sides of head, throat, and sides of neck blackish brown;
under surface generally pale brown, with an admixture of white;
tip of tail white. Total length about 40 cm. (The brownish rather
than blackish tone possibly represents a discoloration that had
developed since the specimen was collected in 1880. A similar dis-
coloration is now observable in the type of L. smithi Rhoads.)
In writing of this specimen, Giglioli says (1881, p. 45) : "The
Natives told Count Marazzani that the Lophiomys is rare, that it
lives in deep holes in the strangely fissured rocks of that country."
He also records a specimen killed at Keren in the Bogos country,
Eritrea, in 1870.
202 EXTINCT AND VANISHING MAMMALS
Oustalet (1902, p. 399) records a specimen from Massaua,
Eritrea.
"This animal is said to occur in the Khor Baraka and also at
Tamai [respectively south and west of Suakin], and it is stated
that it burrows under the roots of trees like a rat" (Anderson, in
Anderson and de Winton, 1902, p. 290) .
All, or nearly all, of the above-mentioned specimens, except the
type of aethiopicus, were recorded as "L. imhausi" before the plas-
ticity of the genus was recognized, but all of the localities are much
nearer to the type locality of aethiopicus than to that of any other
described form.
Imhaus's Maned Rat; Imhaus's Crested Bush Rat
LOPHIOMYS IMHAUSII A. Milne-Edwards
Lophiomys Imhausii A. Milne-Edwards, L'Institut, vol. 35, p. 46, 1867. (Based
upon a specimen secured alive at Aden, Arabia, but of unknown prove-
nance (A. Milne-Edwards, 18676, p. 115) ; Thomas remarks (1910, p. 222)
that Aden is "a place to which Somali animals are very commonly brought
for sale.")
SYNONYM: Lophiomys smithi Rhoads (1896) ("Sheikh Husein, West Somali-
land" = Ethiopia) .
FIGS.: A. Milne-Edwards, 18676, pis. 6-10; Kull, 1894, p. 136, fig.; Rhoads,
1896, pi. 25; A. D. Smith, 1897, p. 64, fig.; Drake-Brockman, 1910, pi.
facing p. 133.
This species is " found probably throughout the Somali country,
but [is] undoubtedly a very rare animal" (Drake-Brockman, 1910,
p. 134).
It is covered with very long silky hairs, of mixed white and black ;
those of the back rising in a crest from the crown to the tip of the
tail, and separated from those of the sides by an area of much shorter
hairs, brittle and grayish tawny; tail long, not prehensile, covered
with hairs like those of the body (A. Milne-Edwards, 1867a, pp. 46-
47) . The general appearance is not ratlike. The dorsal crest is
erectile. An adult male from British Somaliland measured: head
and body, 11 inches; tail, 8 inches. (Drake-Brockman, 1910, p. 133.)
The known distribution includes British Somaliland and south-
eastern Ethiopia.
Kull (1894) describes and figures two specimens from Somaliland.
A specimen (the type of L. smithi) secured by A. D. Smith (1897,
p. 64) at Sheikh Husein, Ethiopia, in 1894 was the only one seen in
a journey of 4,000 miles through British Somaliland, Ethiopia, and
Kenya.
In British Somaliland "I have seen it at Sheikh and near Burao,
but never lower than 4,000 ft. One specimen was killed by Somalis
at Upper Sheikh and one caught alive, while an adult female and
young male were caught near Burao ....
ORDER RODENTIA: RODENTS 203
"Its custom of proceeding with crest erect is in all probability a
protective measure to frighten its enemies, which might mistake
it very easily for a young porcupine." (Drake-Brockman, 1910,
p. 134.)
Goba Maned Rat
LOPHIOMYS BOZASI Oustalet
Lophiomys Bozasi Oustalet, Bull. Mus. Hist. Nat. [Paris], vol. 8, no. 6,
p. 400, 1902. (Goba, southern Ethiopia; alt. 3,000 m.)
This species appears to be known only from some three or four
specimens.
The female type is described as larger than L. imhausii; fur
thicker; an elongate white spot over each eye, with a black band
between; a white spot below the eye; muzzle and area about each
eye black. Total length, 535 mm. (Oustalet, 1902, p. 401.)
De Winton (in Anderson and de Winton, 1902, p. 291) records
(under the name of L. imhausi) a specimen from near "Het Marafia"
( = Let Marefia) and another from the forest of Tikem; both locali-
ties are in Shoa, Ethiopia.
Hollister (1919, p. 37) refers to the present species a specimen
from Let Marefia, Shoa.
Uaragess Maned Rat
LOPHIOMYS THOMASI Heller
Lophiomys thomasi Heller, Smithsonian Misc. Coll., vol. 59, no. 16, p. 4, 1912.
("Mt. Gargues (Uaragess), 6000 feet altitude, Mathews Range, British
East Africa.")
This species appears to be known from only three specimens from
the type locality.
"Allied most closely to ibeanus, differing chiefly in darker and
more contrasting coloration .... General dorsal coloration deep
black, the hairs everywhere broadly white tipped . . . ; the sides
somewhat more extensively white than the median maned area;
. . . lateral bands . . . olive-drab .... Head chiefly black with
two prominent wide white bands over eyes, which meet on forehead,
another large white spot below eye .... Underparts grayish, the
hairs extensively white tipped . . . ; tail silvered like dorsal region,
the extreme tip white. . . . Head and body, 270 mm., tail, 165."
(Heller, 1912, p. 4.)
"These three specimens were caught in rock crevices .... Heller
believes these Lophiomys to be strictly rock-dwellers, notwithstand-
ing reports of their living in holes of trees." (Hollister, 1919, p. 37.)
204 EXTINCT AND VANISHING MAMMALS
Jackson's Maned Rat
LOPHIOMYS TESTUDO Thomas
Lophiomys testudo Thomas, Ann. Mag. Nat. Hist., ser. 7, vol. 15, p. 80, 1905.
("Ravine Station, [Mau Plateau,] British East Africa.")
"The type-specimen . . . remains to this date unique" (Hollister,
1919, p. 37).
This species differs from the others in skull characters; "line of
glandular bristles on sides narrower and less conspicuous than in
other species" (Thomas, 1910, p. 223). "Basal third of underfur
dark brown, middle third white, tip black, the contrast between the
colours more marked than in L. Imhausi. Suborbital white spot well
marked. Dark band dividing the frontal from the auricular white
patch scarcely perceptible. . . . Hairs of lateral line olive. Under
surface hoary grey .... Tail with its underfur mixed whitish and
black, the tip for a length of about half an inch sharply contrasted
white. . . . Head and body 296 mm.; tail 176." (Thomas, 1905,
p. 81.)
Mau Maned Rat
LOPHIOMYS IBEANUS Thomas
Llophiomys] ibeanus Thomas, Ann. Mag. Nat. Hist., ser. 8, vol. 6, p. 223,
1910. ("Mile 513 of the Uganda Railway (between Londiani and Lumbwa
Stations) in Mau region," Kenya.)
SYNONYM?: Lophiomys ibeanus hindei Thomas (1910).
FIGS.: A. B. Baker, 1912, pi. 1; Hollister, 1919, pi. 2.
This is perhaps the least rare species of Lophiomys. It occurs in
the Mau region of Kenya, while the doubtfully distinct L. i. hindei
has been recorded from the neighboring Aberdare Range and from
Mount Kenya (Hollister, 1919, pp. 38-39) .
This species "is coloured like the Abyssinian form referred to
L. bozasi, and has equally prominent lateral stripes." It differs
from other species in skull characters. (Thomas, 1910, pp. 223-224.)
In addition to the type, specimens of L. i. ibeanus are recorded
from El-Burgon and from the Mau Forest near Njoro, Kenya, while
three specimens of L. i. hindei are recorded from the Aberdare
Mountains (Thomas, 1910, p. 224).
A. B. Baker (1912, p. 2) writes:
This species of Lophiomys occurs in the higher part of British East Africa
and is known only to the Wanderobo, a tribe of expert hunters, who explore
every corner of the forests. Mr. Goldfinch was well acquainted both with the
game of that region and with its animals generally, but this one he knew
only from descriptions given by the natives. At his urgent request they secured
two specimens in the forest near Nakuru, at about 8000 feet altitude. . . .
Mr. Goldfinch states that Lophiomys is arboreal and lives in the thick
forest of the high country, . . . also that the natives are averse to handling
ORDER RODENTIA: RODENTS 205
the animal, believing its bite to be poisonous. It is he says, "very rare or only
got by accident here." ... It is strictly nocturnal.
Lonnberg (1912, p. 100) records a specimen from Mau Escarp-
ment.
Hollister (1919, p. 38) records specimens of ibeanus from the
Naivashi Escarpment and from Nakuru, and one of hindei from
Mount Kenya.
"The first one of these animals I got was when I was stationed
at Nakuru; it came from the Aberdare side. It was taken out of a
hole in a tree by a Wanderobo .... I had no difficulty in getting
all I wanted, and at one time I had something like a dozen of them."
(Goldfinch, 1923, p. 1091.)
Family MURIDAE: Old World Rats
The limits of this family, as of the Cricetidae, are not definitely
settled. The two families are similar in the multiplicity of their
genera, species, and subspecies. While the Muridae were originally
confined to the Old World, several forms of Rattus and Mus have
attained world-wide distribution through transoceanic shipping and
are thoroughgoing pests. In the genus Rattus, two species endemic
on Christmas Island in the Indian Ocean have become extinct, and
an Australian subspecies has apparently met the same fate. Single
representatives of two other Australasian genera (Mastacomys and
Zyzomys) are treated in the following pages.
South Australian Spiny-haired Rat
RATTUS CULMORUM AUSTRINUS Thomas
Rattus culmorum austrinus Thomas, Ann. Mag. Nat. Hist., ser. 9, vol. 8,
p. 427, 1921. ("South Australia; type probably from Kangaroo Island."
However, Iredale and Troughton (1934, p. 74) suggest "Port Lincoln" as
the type locality.)
More than a century has elapsed since the type specimen of this
rat was sent to the Zoological Society of London in 1841, and
there seem to be no records of appreciably later specimens.
The fur is sparse, coarse, and more or less admixed with flattened
spines; it is longer in this than in the other subspecies, the hairs of
the back being commonly 20 mm. in length ; general color above gray
rather than fawn color; under parts equally gray. Head and body,
155 mm.; tail, 120 mm. (Thomas, 1921, p. 427; Jones, 1925, pp.
298-299.)
Thomas (1921, p. 427) mentions six specimens besides the type,
and remarks: "Evidently a common rat in South Australia in the
forties, but whether it still exists in any out-of-the-way part of
the colony we have no evidence to show."
206 EXTINCT AND VANISHING MAMMALS
To this Jones adds (1925, p. 299) : "I know of no recent records
or specimens of the species. So far, the out-of-the-way place has
not been found by collectors, and this fact should prove a stimulus
to our field naturalists."
A. S. Le Souef writes (in Hit., February 15, 1937) that this par-
ticular race is probably extinct, but that one or more of the other
subspecies are still numerous at times.
[The other subspecies are: Rattus culmorum culmorum Thomas
and Dollman, of Queensland; R. c. youngi Thomas, of Moreton
Island, Queensland; R. c. vallesius Thomas, of the interior of New
South Wales.]
Captain Maclear's Rat
RATTUS MACLBARI (Thomas)
Mus macleari Thomas, Proc. Zool. Soc. London 1887, p. 513, 1887. (Christmas
Island, eastern Indian Ocean.)
FIGS.: Thomas, op. cit., pi. 42 (colored); Andrews, C. W., 1900, pi. 2 bis,
figs. 1, 3, 6, 7, 8 (skull and teeth).
This rat, isolated on Christmas Island, some 200 miles south of
Java, the nearest land, is believed to have become extinct in the
early years of this century. It is apparently nearest related to
Rattus xanthourus of Celebes and R. everetti of the Philippines,
which it somewhat resembles in appearance.
About the size of a Roof Rat, it is described as grizzled rufous
brown above, the belly but little lighter, pale rufous; longer hairs
black, feet dark like the body. A striking feature is said to be the
prominent long black hairs of the lower back, which, as in the other
related rats, project far beyond the shorter portions of the pelage.
The tail, which equals or slightly exceeds the length of head and
body, is dark in its proximal half, white in its distal portion, and
scaly. The skull is large and strongly built, with beaded supra-
orbital edges, and the anterior edge of the zygomatic plate projects
forward conspicuously. Measurements: head and body, 235-240
mm.; tail, 246-267; hind foot, 48.5-50; ear, 17-17.5; basal length
of skull, 47.5 ; zygomatic width, 26.2. Mammae four.
This island rat was first made known by Thomas (1887) from a
specimen brought from Christmas Island by Captain Maclear of
the British surveying-ship Flying-fish, who procured it on his visit
there in 1886. In the following year additional specimens were
secured by J. J. Lister, who, as naturalist, accompanied a second
expedition to the island on H. M. S. Egeria. At that time the island
was uninhabited and covered with jungle and forest. Of about 40
square miles in area, its highest point is about 1,200 feet above sea
level; geologically, it is largely of coral limestone resting on a basis
ORDER RODENTIA: RODENTS 207
of volcanic rock. The specimens brought back by this expedition
indicated a deposit of phosphate rock, to exploit which a settlement
was founded at Flying-fish Cove, the only anchorage. Shortly after
this, Dr. Andrews made a three-months' visit (in 1897) in order
to survey the natural conditions there. His account contains prac-
tically all that is known of the species, which was then by far the
commonest of the mammals found in the island. He wrote:
In every part I visited it occurred in swarms. During the day nothing
is to be seen of it, but soon after sunset numbers may be seen running
about in all directions, and the whole forest is filled with its peculiar
querulous squeaking and the noise of frequent fights. These animals, like
most of those found in the island, are almost completely devoid of fear, and
in the bush if a lantern be held out they will approach to examine the new
phenomenon. As may be imagined, they are a great nuisance, entering the
tents or shelters, running over the sleepers, and upsetting everything in their
search for food. They seem to eat anything, and destroy any boots or skins
incautiously left within their reach. Their natural food appears to be mainly
fruits and young shoots, and to obtain the former they ascend trees to a
great height. ... In the settlement they utterly destroy all the fruit they
can get at, and frequently come into conflict with the fruit-bats on the
tops of the papaia-trees. A number of dogs is kept to keep them in check,
and near the settlement they are certainly already less numerous than else-
where. In the daytime these rats live in holes among 1 the roots of trees, in
decaying logs, and shallow burrows. They seem to breed all the year round.
After 10 years' absence, Andrews (1909) again visited Christmas
Island for the purpose of ascertaining what changes had taken
place in the interim as a result of white occupation. Such changes
were "chiefly noticeable in the immediate neighbourhood of the
settlement and quarries, while the rest of the island, although tra-
versed by roads in several directions, is practically unchanged." The
rats, however, had gone. For whereas 10 years earlier they were
found everywhere all over the island in abundance, in 1908, in spite
of continual search, not a single specimen of this tree-climbing
species or of the other burrowing rat, R. nativitatis, could be found
in any part of the island. He says further:
This complete disappearance of two such common animals seems to have
taken place within the last five or six years, and to have been the result of some
epidemic disease, possibly caused by a trypanosome, introduced by thfe
ship-rats. These are a variety of Mus rattus, and have been introduced in
considerable numbers, though they do not seem to have spread to the
remoter parts of the island at present, at least to any great extent. The
disappearance therefore of the native forms cannot be due to direct com-
petition with the intruders, but must be the result of disease, a conclusion
supported by an observation made by the medical officer, Dr. McDougal,
who told me that some five or six years ago he frequently saw individuals
of the native species of rats crawling about the paths in the daytime,
apparently in a dying condition.
Since Andrews's second visit in 1908, one or two other zoologists
have visited Christmas Island for the study of its fauna, notably
208 EXTINCT AND VANISHING MAMMALS
M. W. F. Tweedie in 1932, but apparently no one has since found
a trace of the two indigenous rats or of the shrew (Crocidura
fuliginosa trichura) that were abundant before the settlement. No
doubt the supposition that they were exterminated through the
spread of some disease brought in by introduced House Rats is the
most likely explanation of their disappearance. Chasen (1933) , who
has written of the birds of the island, adds that in addition to Rattus
rattus (subsp. ?) , the House Mouse (Mus musculus) and the small
Rattus concolor, a member of a group adaptable to colonization as a
human acolyte, have also been introduced in this island.
G. M. A.
Christmas Island Burrowing Rat; "Bulldog Rat"
RATTUS NATIVITATIS (Thomas)
Mus nativitatis Thomas, Proc. Zool. Soc. London 1888, p. 533, 1889. (Christ-
mas Island, eastern Indian Ocean.)
FIGS.: C. W. Andrews, 1900, pi. 2 (col. fig.) ; pi. 2 bis, figs. 2, 4, 5, 9, 10 (skull
and teeth).
This rat and Rattus macleari (q. v.) are the only indigenous ter-
restrial rodents known from Christmas Island, Indian Ocean, and
are both now believed to be extinct.
Rattus nativitatis was first collected by J. J. Lister in 1887, when
as naturalist aboard H. M. S. Egeria, in the year following the
visit of Captain Maclear, he explored part of the island. On this
occasion, a landing party under Captain Aldrich cut a way through
the jungle to the highest part of the island.
In contrast to the other species, R. macleari, this rat was a more
stoutly built animal of burrowing habits. It is described as a large
species about 17 inches in total length with a tail much shorter than
head and body, of a thickset clumsy form, but having a peculiarly
small and delicate head. In color it was a dark umber brown all
over, the belly not or scarcely paler. The fur of the back, though
long, thick, and coarse, was without the elongated piles characteristic
of R. macleari. The claws were broad and strong, adapted for
digging. Mammae abdominal, three pairs. There is a slight degree
of variation in color, some individuals being a warmer brown than
others, and occasional ones having a small irregular patch of white
fur on the belly. Teeth relatively small and weak. Measurements:
head and body, 275 mm.; tail, 182; hind foot, 50; ear, 24 (these
for the largest of nine specimens). Skull: basal length, 46.8 mm.;
zygomatic width, 24.8; nasals, 20.5; diastema, 15.5; upper cheek
teeth, 7.6. A comparison of the forearm and hand bones in the two
species is given by Forsyth Major (with figures) in Andrews's
(1900) Monograph of Christmas Island.
ORDER RODENTIA: RODENTS 209
This rat apparently was less generally distributed over the island
than R. macleari, inhabiting hilly areas in the interior. Andrews 's
account supplies practically all that is known of it. He wrote:
Though very numerous in places, especially on the hills, e. g. Phosphate
Hill, [it] is very much less common than M. macleari. I never saw one in
Flying Fish Cove [the settlement], though they certainly have been killed
there. They seem to live in small colonies in burrows, often among the
roots of a tree, and occasionally several may be found living in the long,
hollow trunk of a fallen and half-decayed sago-palm (Arenga listeri). The
food consists of wild fruits, young shoots, and, I believe, the bark of some
trees. [It is a] much more sluggish animal than M. macleari, and unlike it,
never climbs trees; and it is difficult to avoid the belief that the former
species is being supplanted by the latter in spite of the abundance of food.
Both animals are strictly nocturnal, and M. nativitatis, when exposed to
bright daylight, seems to be in a half-dazed condition. The Ross family in
Christmas Island have given this species the name "Bull-dog Rat," and this
has been adopted by the Malays.
This was in 1897. When, in 1908, Andrews revisited the island
to see what changes had followed the planting of a settlement there,
he found both species apparently quite gone. "In spite of continual
search, not a single specimen of either species could be found in any
part of the island." This disappearance, as detailed under Rattus
macleari, was conjectured to have taken place about five or six years
earlier, when the medical officer stationed there had frequently seen
individuals of the native rats "crawling about the paths in the day-
time, apparently in a dying condition." Andrews suggests that the
introduced Roof Rat, by then already present in considerable num-
bers, had brought in some epizootic disease to which the native
species had been susceptible, and in consequence they had been
entirely wiped out in the brief space of a few years. (Andrews, 1909,
pp. 101-102.)
At the time of Andrews's first visit he wrote (1900) : "The con-
ditions of life are apparently extremely favourable, food being
always abundant, and the hawk and owl, which are the only possible
enemies [of these rats], feeding mainly on birds and insects. The
consequence of this is that all the species of mammals are extremely
common, and the individuals are always exceedingly fat. Perhaps
Mus [= Rattus] nativitatis, the bull-dog rat as the Cocos Islanders
have named it, is the least numerous, probably because of some
competition with the much more active and versatile M. macleari,
but most specimens of M. nativitatis have a layer of fat from half
to three-quarters of an inch thick over most of the dorsal surface
of the body." Possibly this very abundance of individuals and their
fat condition made them the more susceptible to any disease brought
in from outside.
While conjecture as to the origin of the endemic fauna is more or
less futile, Andrews nevertheless points out that on the whole its
8
210 EXTINCT AND VANISHING MAMMALS
relations are with "Austro-Malayan" rather than with Javan types.
He noticed on several occasions the transport of insects to the island
by storms "which, during the rainy season, blow occasionally from
the northern quarter," but inclines to the supposition that the "rats,
the fruit-bat, and possibly some of the land birds, very probably
owe their introduction to the island" to the transport by rafts of
trees brought by the equatorial drift from the Timor Sea. This
island is at least of unusual interest as affording a case in which
the native fauna has within a few years been altered as a result of
settlement by man, and two of its few native mammals have be-
come extirpated.
G. M. A.
Broad-toothed Rat
MASTACOMYS FUSCUS Thomas
Mastacomys juscus Thomas, Ann. Mag. Nat. Hist., ser. 5, vol. 9, p. 413,
1882. ("Tasmania.")
This rare rat survives in the Otway Forest of Victoria and in
Cradle Valley, Tasmania, but little or nothing is known of its present
occurrence elsewhere.
Fur long and soft; general color dark grayish brown above and
below, the dorsal hairs tipped with light brown and the ventral
hairs with white; tail and feet dark brown. Head and body, 142
mm.; tail, 95 mm. (Thomas, 1882, pp. 413-415.) It is a large rat,
with a stout build, strong limbs, and a short tail. Head and body
(Tasmanian specimens), 170-182 mm.; tail, 110-113 mm. (Finlay-
son, 1933a, pp. 126, 128.) Victorian specimens have longer tails
(119-124 mm.) (Brazenor, 1934, p. 161).
The type specimen, from an unspecified locality in Tasmania, was
acquired by the British Museum in 1852, and for 80 years no further
information seemed to be forthcoming as to its occurrence in that
state. In 1931, however, five specimens were collected in Cradle
Valley, northwestern Tasmania, at an altitude of about 3,000 feet.
The Broad-toothed Rats were living in colonies in grassy areas on
open heaths, in association with Eastern Swamp Rats (Rattus
lutreolus). "Both rats are probably quite numerous, but the laby-
rinths are the chosen hunting grounds of Dasyurus vivverinus [sic],
and it was not until several days trapping had got rid of the latter
that rats began to be caught." (Finlayson, 1933a, pp. 125-126.)
Doubtless this Native Cat acts as a check upon the increase of
the rats.
Lydekker (1885, p. 227) records some bone fragments of Masta-
comys juscus from the caves of the Wellington Valley, New South
Wales.
ORDER CARNIVORA: CARNIVORES 211
Brazenor (1934, pp. 159-160) records specimens from the follow-
ing localities in Victoria: Swan Island; Gippsland; Layer's Hill
in the Otway Forest; Olangolah, near Beech Forest, at the head
of the Gellibrand River. He also mentions a specimen, apparently
previously overlooked, from the "West Coast of Tasmania, 1872."
He writes (in litt., March 3, 1937) that the species "still survives
in the Otway Forest but not in any numbers."
While-tailed Rat
ZYZOMYS ARGURUS ARGURUS (Thomas)
Mus argwrus Thomas, Ann. Mag. Nat. Hist., ser. 6, vol. 3, p. 433, 1889.
("South Australia.")
This South Australian rat is apparently known only from the
type specimen.
The fur is crisp; general color above pale sandy rufous; ears
rounded, thinly covered with fine white hairs; muzzle and under
parts white, the line of demarcation on the sides not sharply defined ;
hands and feet pure white ; tail uniform white above and below, the
tip slightly penciled. Head and body, 83 mm.; tail, 101. (Thomas,
1889, pp. 433-435.)
"Of this very distinct species there is no material available in
South Australia, which was the home of the type specimen. . . .
"There seem to be no recent records of this remarkable little
rat, and no observations on its habits. Probably it is one of the
many lost species of which no specimens are preserved in our State
collections." (Jones, 1925, pp. 336-337.)
A. S. Le Souef remarks (in litt., February 15, 1937) that these
native rats do not stand up to settlement or invasion of their habitat
by Rattus rattus.
E. Le G. Troughton writes (in litt., April 16, 1937) that the lack
of records since the original description in 1889 supports Wood
Jones's conclusion that the species may be lost from the state.
[According to L. Glauert (in litt., March 17, 1937) , the Western
Australian subspecies, Z. a. indutus (Thomas), is "not reduced in
numbers."]
Order CARNIVORA: Carnivores
Family CANIDAE: Wolves and Foxes
The Canidae are nearly cosmopolitan, indigenous species being
found in all important land masses except Madagascar, the central
and eastern parts of the Malay Archipelago, New Zealand, and
212 EXTINCT AND VANISHING MAMMALS
Tasmania. There are about 19 'genera and more than 200 species and
subspecies. The predatory habits of some of the larger species
(especially the wolves) bring them into conflict with the economic
interests of man, and the animals have suffered accordingly par-
ticularly in North America. Accounts of no less than 24 New World
forms appear in Dr. Allen's volume (1942), while only 4 Old World
forms are treated herein. One of the these, the Japanese Wolf, is
extinct.
Abyssinian Wolf; Abyssinian Red Wolf. Cuberow (Ethiopian)
SlMENIA SIMENSIS SIMENSIS (Ruppell)
Canis simensis Ruppell, Neue Wirbelthiere zu der Fauna von Abyssinien
gehorig, Saugethiere, p. 39, 1835. (Mountains of Simien, Abyssinia.)
FIGS.: Ruppell, 1835, pi. 14; Mivart, 1890, pi. 6; Bryden, 1899, pi. 15, fig. 8;
Lydekker, 1908, pi. 15, fig. 8; Fuertes, Abyssinian Birds and Mammals,
pi. 29, 1930.
This interesting animal has a restricted range and occurs in
limited numbers. The Committee of Experts of the Second Inter-
national Conference, held at London in 1938, states (1938, p. 8) that
this species is "almost completely confined to Abyssinia," and sug-
gests its inclusion in Class A of the Annex at the next Conference
for the Protection of African Fauna and Flora.
Snout long and slender; general color light yellowish reddish
brown, mixed with black on the sides; white about the mouth, eyes,
inner margins of ears, chest, belly, lower parts of limbs, and lower
side of tail toward base ; distal half of tail blackish, and upper side
toward base mixed with black. Head and body, 99 cm. ; tail, 25 cm.
(Mivart, 1890, pp. 18-19.)
"We observed this wolflike dog in the mountains of Simien,
where it lives in packs, and hunts tame sheep and small game, but
never becomes dangerous to man. It occurs also in most of the
other Abyssinian provinces. Its vernacular name in Simien was
given to me as 'Kaberu.' ' (Ruppell, 1835, p. 39, transl.)
"Since Riippell's time little has been heard of this wolf and scarcely
any fresh or recent information is to be obtained concerning it. From
its predatory habits it is probable that the Abyssinians, so soon as
they began to acquire fire-arms, turned their attention to its destruc-
tion, and that in consequence it has become much scarcer than it
used to be. ...
"It would be extremely interesting to know if this handsome wolf
still survives -in Abyssinia in any numbers. Modern travellers and
sportsmen apparently make no mention of it." (Bryden, 1899,
pp. 601-602.)
Lydekker (1908, p. 462) refers to "its rarity and zoological in-
ORDER CARNIVORA: CARNIVORES 213
terest." The Cuberow "was scarcely known in England, except by
its skull, till a few years ago, when skins were brought home by
Major Powell-Cotton." The latter saw several of the animals alive
in the mountains of Simien.
Maydon (1932, pp. 220-221) writes that "the Red Wolf is com-
mon" at Simien, and refers to it as being seen occasionally on the
Gojam plateau, between Lake Tsana and Addis Ababa.
W. H. Osgood (oral communication, 1936) speaks of this species
as not uncommon locally. The Field Museum expedition of 1926-27
obtained about five specimens. Alfred M. Bailey (oral communi-
cation, 1937) does not consider that it is in any danger from the
Ethiopians.
This animal appears to occupy a peculiar zoological position.
Lydekker considers it neither a wolf nor a jackal, while Pocock
denies to it affinity with the foxes.
[A subspecies from south-central Ethiopia has been proposed by
De Beaux under the name of Cam's (Simenia) simensis citernii (Atti
Soc. Ital. Sci. Nat., vol. 61, p. 25, 1922; type locality, "Arussi:
Barofa"). No information is at hand concerning its numerical
status.]
Japanese Wolf
CANIS HODOPHILAX Temminck
Canis hodophilax Temminck, Tijdschr. Natuurl. Geschied. Physiol., pt. 5, p. 284,
1839. (Japan; i. e., Hondo.) (Cj. Harper, 1940, p. 192.)
FIGS.: Temminck, 1842-45, pi. 9; Mivart, 1890, p. 14, fig. 17; Beddard, 1902,
p. 418, fig. 209.
This wolf is now considered extinct.
It is distinguished from the European Wolf by its smaller size
and shorter legs, though it differs but little in the nature and color
of its pelage; fur short and smooth, but tail bushy; ground color
gray or ashy; basal two-thirds of the hairs of back and rump thus
colored, the tips black ; sides, neck, belly, and tail gray, the extreme
tips of the hairs blackish; head and muzzle dark gray; lips more
or less whitish ; outer surface of ears brownish rufous ; four extremi-
ties gray, washed with rufous and brown; tail tip without colored
tuft. Height at shoulder, 16 inches; total length, 3 feet 9 inches, of
which the tail comprises about 1 foot; ears, 3 inches. (Temminck,
1844, pp. 38-39.) "Prof. Brauns . . . says that in the Museum at
Tokio there are very differently coloured skins, namely 'yellowish/
'brownish,' and 'whitish grey' " (Mivart, 1890, pp. 14-15).
The Japanese Wolf lives in wooded and mountainous regions, and
hunts in small family parties. It is as much dreaded by the Japanese
214 EXTINCT AND VANISHING MAMMALS
as the European Wolf is in its range. It shows itself often in winter,
notwithstanding the assiduous pursuit of which it is the object. The
Japanese state that its flesh is unwholesome. (Temminck, 1844,
p. 39.)
A female wolf from Japan was presented to the Zoological
Society of London in 1878 (Flower, 1929, p. 114) . Mivart (1890,
p. 15) records a skull in the British Museum from the province of
Kotsuke.
Thomas (1906, p. 342) records a specimen collected in 1904 or
1905 in the vicinity of Washikaguchi, Nara Ken, Hondo. The col-
lector, M. P. Anderson, adds: "The Wolf was purchased in the flesh,
and I can learn but little about it. It is rare, some say almost
extinct."
Aoki (1913, p. 317) gives the range of this animal as "Hondo
(Thomas), China." Hatta remarks (1928, p. 1033): "Cam's hodo-
phylax T. confined in Japan to the heart of Hondo, Yamato and
Wakayama, occurs also in China." These reports from China are
considered erroneous. Pocock (1935, p. 658) records a skull from
Chichibu.
Nagamichi Kuroda writes (in litt., July 5, 1938) that many of
these wolves were formerly said to be seen in the mountainous dis-
tricts of Hondo, but that the animal is now considered completely
extinct. It was destroyed because of its injuriousness to men and
cattle. It is said that the only specimens in Japan are a mounted
male from Fukushima Prefecture, Hondo, which is now preserved
in the Tokyo Science Museum, and one or two skulls.
Kuroda (1938, p. 36) records the animal from the following addi-
tional localities in Hondo: Rikuchu, Shimotsuke, and Aomori.
Yezo Wolf
CANIS LUPUS HATTAI Kishida
Cam's lupus hattai Kishida, Lansania, vol. 3, no. 25, p. 73, 1931. (Sapporo,
Hokkaido, Japan.)
SYNONYM: Cants lupus rex Pocock (1935).
Although extinct in Hokkaido (or Yezo), this wolf survives in
Sakhalin and perhaps in the Kuriles.
It is much larger than Cam's hodophilax of Japan and is dis-
tinguished from C. I. lupus of Europe by its larger premolar teeth
and by its longer palate and mandible (Pocock, 19356, p. 659).
In the Amur region, according to Schrenck (1859, pp. 45-48),
the wolf is most numerous in northern Sakhalin. Its principal object
of chase is the wild Reindeer. Occasionally packs approach the
villages or solitary houses of the natives and destroy their dogs. The
animal ranges to the south end of Sakhalin and occurs also on the
Kuriles.
ORDER CARNIVORA: CARNIVORES 215
Pocock (1935b, pp. 659-660) refers to "the discovery of this big
wolf in Yeso in the early 'eighties/' and to the opinion of Brauns
(1881) "that it possibly inhabited the Japanese islands . . . between
Yeso and Kamschatka."
"Canis lupus L. is found ... in Sakhalin; in Hokkaido it was
abundant some thirty years ago, but it has decreased so that it seems
to be totally exterminated at present" (Hatta, 1928, p. 1037) .
"Aoki and Kishida both reported it from this island [Sakhalin]
and Hokkaido (rare) " (Kuroda, 1928, p. 226) .
"In authentic historic times the wolves occurred in the main
island of Hokkaido, in Sakhalin, and in Kunashiri, Etoruf and
Paramushir of the Kurile Islands. It seems true that the wolves
were not so frequent in Hokkaido as compared with the other
mammals. They were still fewer in Sakhalin and in the Kurile
group. Though old records say that the wolves fed mostly upon the
deer which abounded in Hokkaido, at the beginning of settlement
they wrought serious havoc amongst herds. So the government at
that time paid a high bounty for the slaughter of the animal. For
instance, the local government in Sapporo paid 7 yen for one wolf
from 1878 to 1882 and 10 yen for each from 1883 to 1885. More
than 1500 wolves were brought in for the bounty during the 11 years
from 1878 until 1888. Since then we have heard scarcely any account
of the animal in Hokkaido." (Inukai, 19326, p. 525.)
Kuroda (1938, p. 36) gives the range of this subspecies as Sak-
halin, the Kuriles, and Hokkaido ; on this last island it is extinct.
Japanese Raccoon-dog
NYCTEREUTES PROCYONOIDES VIVERRINUS (Temminck)
Canis viverrinus Temminck, Tijdschr. Natuurl. Geschied. Physiol., pt. 5,
p. 285, 1839. (Japan.)
FIGS.: Temminck, 1842-45, pi. 8; Martens, 1876, pi. 1.
Formerly abundant in Japan, this animal has become extremely
scarce.
The form is small and foxlike; the tail is short and bushy. The
general color is yellowish brown; hairs of the back, shoulder, and
tail tipped with black; arms and legs blackish brown; a large dark
brown spot on each side of the face, beneath and behind the eye
(Martens, 1876, p. 78). The measurements of some representative
of the species on the Asiatic mainland are given by Mivart (1890,
p. 135) as follows: head and body, 530 mm.; tail, 140 mm.
A century ago the Raccoon-dog was considered very common in
Japan (Temminck, 1844, p. 40) . At this period "Siebold found it to
be very common throughout the Japanese islands, where its flesh
was considered as good food with an agreeable flavour, and its
216 EXTINCT AND VANISHING MAMMALS
powdered, calcined bones a valuable medicine. ... It is not deemed
destructive to poultry. The natives employ its skin to make bellows,
and also to decorate their drums and for winter head-gear." (Mi-
vart, 1890, p. 135.)
Possibly Ognev refers to its former rather than to its present
status when he writes (1931, p. 369) that it is widely distributed in
Japan and particularly common on Hondo (Honshiu) Island.
In the open season extending from October 15, 1929, to April 15,
1930, 15,218 of these animals were taken in Japan (Uchida, 1935,
p. 8.)
FIG. 22. Japanese Raccoon-dog (Nyctereutes procyonoides viverrinus}
"Muko-jima, a small island in the Inland Sea on the coast of
Yamaguchi Prefecture, is famous as a sanctuary for the raccoon dog
.... In feudal times it was abundant throughout this country.
Owing to indiscriminate hunting it became extremely scarce." (Ka-
buraki, 1934, pp. 4183-4184.)
Uchida writes (1935, p. 25) that the animal is gradually becoming
scarce in Japan, since its fur is highly valued and a large number
of skins are exported annually. Mukojima is inhabited by innumer-
able individuals. It is supposed that persecution on the Japanese
mainland gradually forced them to migrate to the island, where
they found a safe breeding place and an abundance of fish for food.
[Concerning the several mainland representatives of this species,
Arthur de C. Sowerby writes (in litt., April 24, 1937) :
"There are now several subspecies of the raccoon-dog recognized in
China and neighbouring regions, namely, Nyctereutes procyonoides
ORDER CARNIVORA: CARNIVORES 217
procyonoides (Gray) of Central and South China, Nyctereutes pro-
cyonoides orestes Thomas of South-west China, and Nyctereutes
procyonoides ussuriensis Matschie of North-east China, Manchuria,
and neighbouring eastern Siberia and Korea. Except in a few places
these animals are probably as plentiful to-day as they ever were,
except that they seem to have been badly decimated in the Amur
region, where once they were particularly plentiful, for the sake of
their pelts. In years past large quantities of the latter have been
exported from China, but at present there is no demand on either the
American or European markets for the 'raccoon' or 'raccoon-dog'
skins, as they are known to the trade, and consequently none are
coming from the interior to the ports, except such as are required
for the home market, which is fairly considerable."
A few years ago approximately one-half million skins were ex-
ported annually from Shanghai (Sowerby, 1934a, p. 287).]
Family URSIDAE: Bears
The bears occur on all continents except Australia and perhaps
Africa (where a single problematical, extinct species has been re-
ported from Morocco and Algeria). The only South American
species is restricted to the Andean region. Thus the distribution of
the Ursidae is not quite so extensive as that of the Canidae. The
bears have suffered perhaps even more than the wolves at the hands
of man. Seven genera and about 135 forms are provisionally recog-
nized. The majority of the latter, however, are North American
Grizzly Bears, whose exact taxonomic status remains somewhat
uncertain. Dr. Glover M. Allen, in his volume on New World mam-
mals (1942), treats all the Grizzly Bears and some of the Black
Bears, as well as the South American Spectacled Bear. The present
volume deals with the various forms of the Brown Bear in Europe
and Asia and with the Atlas Bear of North Africa.
Old World Brown Bear. Ours brun (Fr.) . Brauner Bar (Ger.) .
Oso (Sp.). Orso bruno (It.)
URSUS ARCTOS Linnaeus
[Ursus] arctos Linnaeus, Syst. Nat., ed. 10* vol. 1, p. 47, 1758. (Sweden.)
FIGS.: Geoffroy and Cuvier, Hist. Nat. Mamm., vol. 5, pi. 211 (U. a. pyrenai-
cus), pi. 212 (U. a. collaris), 1824; Gervais, Hist. Nat. Mammif., pt. 2,
pi. 13, 1855 (U. a. pyrenaicus) ; Fitzinger, Bild.- Atlas, Saugth., fig. 72,
1860; Millais, 1904, pi. facing p. 236; Martin, 1910, pi. 38; Cabrera, 1914,
pi. 5 (U. a. pyrenaicus} ; Ognev, 1931, pi. 1 (U. a. caucasicus) ; Pocock,
1932, pi. 2, upper fig.; Castelli, 1935, pis. 1, 2.
Numerous forms of the Brown Bear have been described, but
there is no general agreement on the validity of most of them, and
218 EXTINCT AND VANISHING MAMMALS
the systematics of the group remains in a state of very considerable
confusion (cf. Miller, 1912, pp. 285, 296; Ognev, 1931, pp. 14-118;
Pocock, 1932). In the account that follows, only two of the forms
will call for separate treatment. Among those that have been de-
scribed, the following may be mentioned as being better known or
as having gained more or less recognition in the literature:
Ursus arctos arctos Linnaeus, of northern Europe.
U. a. pyrenaicus Fischer, of the Pyrenees and northern Spain.
U. a. meridionalis Middendorff, of the Caucasus.
U. a. syriacus Hemprich and Ehrenberg, of Asia Minor.
U. a. collaris Cuvier, of Siberia.
U. a. beringianus Middendorff, ranging from Manchuria to Kam-
chatka.
U. a. lasiotus Gray, of Mongolia, Manchuria, Hokkaido, and the
Kuriles.
U. a. isabellinus Horsfield, of the western Himalayas and the
Thian Shan.
U. a. pruinosus Blyth, of Tibet (not generally regarded as con-
specific with U. arctos, but so treated by Pocock, 1932).
The Brown Bear has become extinct over the greater part of its
former range in western Europe but survives in small numbers in
remote and chiefly mountainous areas in Norway, Sweden, Spain,
France, Austria, Czechoslovakia, Italy, Yugoslavia, Greece, Bul-
garia, Rumania, Poland, and Estonia. It has remained much com-
moner in parts of Russia and northern Asia.
The general form of the species is short and heavy; fur long and
rather loose; head moderately pointed, broad posteriorly; ear short
and rounded; front claws strongly curved, blunt, at least twice as
long as hind claws ; tail very short, concealed in the fur. The general
color is usually a light brown or dull buff, the head not essentially
different, but feet and outer surface of legs darker. There are many
individual and racial variations in color. Measurements of an adult
male from Sweden: head and body, 1,900 mm.; tail, 80 mm.; hind
foot, 195 mm.; ear, 90 mm. (Miller, 1912, pp. 287-296.)
The range of the species is the "entire continent of Europe wher-
ever sufficiently extensive forests remain; east into Asia" (Miller,
1912, p. 287).
Great Britain. Numerous postglacial remains have been found
in various parts of England. These include bones from refuse heaps
that are probably of Roman origin. The remains found in Ireland
appear to belong to an older species than Ursiis arctos. In ancient
times in Britain the animal was trailed with boar-hounds and at-
tacked with arrows, pikes, clubs, javelins, and long knives. The
great Caledonian forest in Scotland seems to have been the chief
ORDER CARNIVORA: CARNIVORES 219
stronghold of the British bears. Bears were transported from Britain
to Rome, probably in the fourth century. After the extinction of
the species in Britain, foreign animals were imported for the pur-
pose of "bear-baiting." This was done in the reigns of Henry II,
Henry VIII, Queen Elizabeth, and Queen Anne. Queen Elizabeth
entertained the French Ambassador and the Danish Ambassador
at different times with such a spectacle. (Harting, 1880, pp. 11-29.)
Many place names in Wales afford evidence of the former occur-
rence of bears. Boyd Dawkins thinks that they became extinct in
Britain before the tenth century. Bell, in his British Quadrupeds,
says that they may have existed in Scotland as late as 1073, but
tradition gives the latest date as 1057. (Millais, 1904, pp. 236-237.)
Norway. According to Collett, the bear was numerous about
1750 and did great damage to the cattle all over the country, but
during the following 150 years the numbers were reduced owing to
improved firearms and to high rewards paid for animals shot (Hj.
Broch, in litt., December, 1936).
Bowden states (1869, p. 4) that "the Brown Bear ... is pretty
common in all parts of this country, especially in Nordland and the
central districts."
"The average number that is killed yearly amounts to ... 250"
(Barnard, 1871, p. 262).
"The number of bears in Norway is now beyond doubt very small.
... A conservative estimate would be that there are not more
than 20-25 bears in Norway south of the Trondhjem Fjord. In the
northern parts the number is beyond doubt also very small, chiefly
consisting of stray bears from Sweden and Finland. . . .
"We have done everything in our power to attempt a protection
sufficient to stop the extinction which obviously threatens the species
in this country. But all attempts have been without avail, as it
cannot be denied that the bears occasionally do some damage to
sheep ....
"We have attempted the establishment of a reservation in certain
forest tracts owned partly by private owners and partly by the
government. But the project has until now failed." (Harald Platou,
in Hit., November 22, 1932.)
By a law of 1932 the bear may be killed only from May 15
to November 1. Previously "the bear, being considered a pest, re-
ceived no protection at all. In fact there was a premium for killing
him ....
"This recent protection ... is due to the efforts of the Norwegian
Association for Hunting and Fishing, which endeavored to induce
the Storting to protect the bear all the year round. The Association
had even collected money which it offered to place at the disposal
of the Norwegian Department of Agriculture to compensate for
220 EXTINCT AND VANISHING MAMMALS
losses to farmers and goat and sheep owners for damage done by
the bear. . . . This offer was not accepted. . . .
"It is not known exactly how many bears there are in Norway.
. . . The greatest number are to be found in the tract Eastern
Hallingdal the Hemsedal mountains to Laerdal, where it is esti-
mated that there are from 15 to 25 bears. It is possible that there
are a few bears between Valdres and Gudbrandsal and in the central
part of Telemarken, but their number is uncertain. There are also
a few bears in the northern part of Norway which come from Swe-
den." (Julius Wadsworth, in litt., July 28, 1932.)
"An area of about 125,000 acres north of the city of Lillehammer
has been privately set aside for the protection of the bear." Mr.
Platou "thinks the bears will now be preserved from extinction."
(Julius Wadsworth, in litt., May 9, 1933, and July 20, 1933) .
Sweden. The bear was formerly found in all parts of the coun-
try, but in most of the provinces constituting Gotaland it had prac-
tically disappeared during the eighteenth century. ^ It has been
calculated that in the whole country 1,351 bears were killed during
the period 1827-1836, and 1,055 during the period 1847-1856. In
the 50-year period from 1856 to 1905, 2,762 bears were killed in
Sweden, including 86 in Wermland, 499 in Dalecarlia, 171 in Gavle-
borg, 144 in Westernorrland, 796 in Jemtland, 292 in Westerbotten,
and 770 in Norrbotten. The rapid decrease is illustrated by the
following statistics on the numbers killed in six of the above-
mentioned districts: 908 in 1856-1865; 434 in 1876-1885; 109 in
1896-1905. In olden times a small bounty was paid on each bear
killed, and in 1864 the amount was raised to 50 riksd. Sportsmen
and others objected to the bounty, and it was finally abolished in
1893. By that time the bears were greatly decimated and in most
provinces entirely exterminated. The building of railroads had con-
tributed decidedly to this decrease. After considerable agitation
for protection of the species, a new law of 1912 declared that no
bear could be shot on crown land without special permission from
the King, unless it had attacked man or domestic animals, nor on
private land without the permission of the owner. According to a
law of 1927, a bear killed anywhere belongs to the Crown. Com-
pensation for damage by bears is now provided by the state. The
whole sum thus paid for domestic animals (sheep, goats, reindeer,
and one horse) , during 1933 and 1934 did not amount to more than
2,404 kr. in all. Since the bears have been protected their numbers
have increased only a little.
In considering the bear's economic status, it may be noted that
it never attacks man unless directly provoked or wounded. It preys
on domestic animals only exceptionally, and not regularly. Many
reindeer succumb to starvation and disease, and when the bear feeds
ORDER CARNIVORA: CARNIVORES 221
on their carcasses, it is often accused of having killed the animals.
Its diet consists largely of insects and their larvae and various kinds
of plant food, especially berries. It also catches voles and lemmings.
(Einar Lb'nnberg, in litt., October, 1936.)
Under present conditions the bears are not threatened with ex-
tinction (Einar Lonnberg, in litt., November 15, 1932) .
Spain (U. a. pyrenaicus) . Cabrera (1914, p. 153) gives the range
of this race as the Pyrenean and Cantabrian districts: Pyrenees
of Aragon and Catalonia; mountains of Santander and Asturias;
extreme north of the Provinces of Palencia and Leon and the
eastern part of Lugo. In historic and even comparatively recent
times it ranged more to the south, reaching at least the center of
the peninsula. In 1582 Argote de Molina reported it not far from
Madrid.
"Bears still occur not unfrequently all along the Cantabrian
range of mountains. On the central chain of Spanish mountains
they seem to be rarer. There are none now in Portugal. Formerly,
as lately as the sixteenth century, before the devastation of the
forests, the bear seems to have had a much "wider distribution in
the Peninsula." (Gadow, 1897, p. 362.)
In Asturias it nightly raids the maize-fields in the valleys in
September. It is also in the habit of attacking and destroying many
cattle. It is tracked to its covert, and a drive with beaters is organ-
ized. From 20 to 30 bears are killed in Asturias every year.
(Marquis de Villaviciosa de Asturias, in Chapman and Buck, 1910,
pp. 296-297.)
France (U. a. pyrenaicus in the Pyrenees; U. a. arctos in the
Alps). Trouessart states (1884, pp. 195-196) that the species is
restricted to the forested and the wildest regions of the Alps and
the Pyrenees. It occasionally ravages the wheatfields and the vine-
yards. It becomes more carnivorous with age and then forms the
habit of making raids upon sheep and calves, and finally it even
attacks grown cattle and horses.
E. Bourdelle (1937, pp. 178-181, and in litt., March 6, 1937) gives
the following account:
Formerly rather widely spread in the mountainous regions of
France Vosges, Jura, Cevennes, Alps, and Pyrenees it disappeared
from the first three areas during the past century, and it now exists
only in the Alps and the Pyrenees. It is generally believed to have
disappeared from the French Alps and that the last two animals
were killed in 1898 in the Forest of Vercors in these mountains.
However, fresh tracks were observed in the same region in 1913, in
1928, and again during the past few weeks. The extent and wildness
of the Forest of Vercors militate in favor of the possibility of a
few bears surviving there.
222 EXTINCT AND VANISHING MAMMALS
In the Pyrenees the Brown Bear, though much less abundant
than formerly, is still met with in the wild areas of upper Ariege
adjacent to Andorra, in the Hautes Pyrenees (massif of Maladetta
and Cirque de Gavarnie), and as far as the neighboring parts of
the Basses Pyrenees (Forest of Irruti, for example). It is even
probable that there are some bears in other parts. While it was
still rather common 20 or 30 years ago, it has been gradually pushed
back into refuges more and more restricted, where it is easily hunted,
so that it has become rarer and rarer during recent years.
Its only economic importance in the Pyrenees consists in its being
a true game animal, its flesh being much prized in the whole region.
Its hide provides a good fur, but not a very valuable one.
The bear of the Pyrenees, like that of the Alps, is the victim
of man's increasing penetration into the mountains, of the extension
of agriculture, forestry, and mining, of highways, railroads, tourist
traffic, and especially the sport associated with the hunting of this
animal.
Protection should be provided for the Pyrenean bears as well as
for those of the Alps (if the latter still exist in the Forest of Vercors) .
Prohibition of hunting and of the sale of flesh and hides would
suffice to halt the steady depletion and perhaps to assure the preser-
vation of the species.
According to Didier and Rode (1935, p. 268) , the last bears in the
Hautes-Alpes were killed at the beginning of the nineteenth century.
Two were killed in 1896 in the massif between La Chambre and
Saint-Remy (Savoie) and Allevard (Isere).
Belgium and The Netherlands. The species no longer exists in
these countries (Martin, 1910, p. 38a; Ognev, 1931, p. 40).
Germany. Blasius (1857, p. 199) reports the bear as still occur-
ring in the Bavarian highlands.
The last specimens were killed in 1759 in Thuringia, in 1770 in
Upper Silesia, in 1810 in the Bavarian-Bohemian forests, and in
1835 in Frauenstein, Bavaria. In the sixteenth century the species
was still of frequent occurrence in Germany. When improved fire-
arms were introduced, the population started a real war against the
animal. The hide was readily sold, and bear-hunting, which in-
volved some danger, was considered by the upper classes as enter-
tainment and sport. Bounties were often paid, as the animals were
looked upon in many places as seriously injurious to cattle. (Inter-
nationale Gesellschaft zur Erhaltung des Wisents, in litt., October,
1936.)
The last bear in Pomerania was killed in 1750; in the Riesenge-
birge about 1800. The species was still being taken in East Prussia
up to 1806. (Krumbiegel, 1930, p. 6.)
ORDER CARNIVORA: CARNIVORES 223
Castelli (1935, p. 33) quotes the Alpines Handbuch (1931) to the
effect that the last one was killed in Wetterstein, Bavaria, in 1864.
Denmark. The bear was once generally distributed over Den-
mark. Remains have been found in the ancient kitchen middens,
but there is no information on the occurrence of the species within
historic times. (Winge, 1908, p. 127.)
Switzerland. In 1869 Fatio (pp. 301-302) gave the following
account of the bear's status: It was formerly abundant in the
north and center of Switzerland but has gradually retired to the
high Alps. It is now scarcely found except in Orisons, in Tessin
(where 9 specimens were killed from 1852 to 1862), and here and
there in the Jura. It has almost entirely disappeared in Valais
and Uri. Basle, Lucerne, Schwyz, and Berne have no more bears.
Castelli (1935, pp. 25-27) supplies the following records: The last
bears near Zurich were killed or recorded in 1565, in Unterwald in
1664, in Fribourg in 1698, while in Solothurn 38 were killed from
1507 to 1737. Other last records are: Berne, 1815; Glarus, 1816;
Vaud, 1843; Valais, 1860; Uri, 1898. In the Engadine 5 were killed
in 1852, 8 in 1861, 6 in 1872, and 4 in 1873. In Orisons 25 were
killed from 1878 to 1887, 9 from 1888 to 1897, and 3 within the
following decade; the last one was killed in 1904 in Val Minger, but
a female with two cubs was reported seen as late as 1919 in Val
Lavirum.
The species is now of exceptional occurrence in Switzerland, being
represented only by an occasional straggler across the border from
western Trentino, Italy (Tratz, in Castelli, 1935, p. 9).
Italy. The bear is now restricted to two general areas in Italy
the extreme north and the mountains of Abruzzi.
From 1837 to 1852 146 specimens were killed in Trentino. The
species is now protected there by the Italian Government. (G.
Schlesinger, in Hit., March, 1937.)
According to Castelli (1935, pp. 50-135), 77 bears were killed in
Trentino from 1886 to 1912. In the district of Cles, at the north
end of the Group of Brenta, the following numbers were killed: 26
from 1886 to 1891; 5 in 1895; 2 in 1900; 1 in 1901; 2 in 1902; 4 in
1903; 1 in 1906; 2 in 1908; 2 in 1909; 3 in 1910; 8 in 1911; 2 in
1912; 3 in 1913. In Trentino 15 bears have been killed from 1922
to 1933, and a small number have been seen yearly up to 1935. The
Brenta Group and vicinity form the last refuge of the species in
Trentino. It is sedentary there, and is in urgent need of protection,
such as would be afforded by the establishment of a National Park.
Castelli (1935, p. 28) quotes Cermenati to the effect that 40 bears
were killed between 1876 and 1886 in Valtellina, Lombardy, Italy.
He adds (p. 31) that the last individuals were killed in Valtellina in
1896 and 1902. He also mentions (p. 32) a report of Depoli in 1928
224 EXTINCT AND VANISHING MAMMALS
that there were about 10 bears, protected by law, in the Province of
Carnaro, northeastern Italy.
In the National Park of the Abruzzi about 200 Brown Bears are
well protected (Tratz, in Castelli, 1935, p. 9). They are found in
the mountains about the valley of the Sangro, and must be regarded
as indigenous, notwithstanding the local tradition that the Czar of
Russia had sent King Ferdinand of Naples a couple of such animals,
which he set free in the mountains of the Abruzzi (Colosi, 1933,
pp. 48-49). The park administration estimates the present number
at about 100 (Laboratorio di Zoologia Applicata a Caccia, in Hit.,
September, 1936) .
The bear is completely gone from the Sila Montains, Calabria,
though present there in the middle of the last century (Hecht, 1932,
p. 23).
Austria. This was probably an indigenous species all over Aus-
tria in former days. In Carinthia it was generally distributed up to
1850; one bear was killed during each of the years 1895, 1920, 1927,
and 1936. They are supposed to have come from the reserves in
Gottschee, Carniola, and on Schneeberg (Monte Nevoso), north of
Fiume, Italy ; perhaps also from Croatia. In Lower Austria the bear
was observed rather frequently up to the last half of the nineteenth
century ; here, in Semmering, Schneeberg, Rax, and the mountainous
areas to the westward, fine stocks of bears were to be found. The
last one was observed in 1919 near Rohr in the mountains of Lillien-
feld. In Upper Austria and Salzburg the species was probably quite
common up to the middle of the nineteenth century. In Tyrol the
decrease started in 1570. At that time Duke Albrecht prohibited
the capture and killing of bears. During the Thirty Years' War the
numbers increased again. Up to about 1840 the annual kill was
from 20 to 30 specimens. The last one was shot in Stellental, Tyrol,
in 1898; in Vorarlberg, in 1870. The bear is not compatible with
cattle-raising or with the increase in human population. (G. Schles-
inger, in Hit., March, 1937.)
According to the Alpines Handbuch (1931), 34 bears were killed
in Tyrol in 1835, and in the same year the last one was killed in the
Schneeberg district near Vienna. The last one was seen at Kar-
wendel, on the Tyrolean-Bavarian border north of Innsbruck, in
1896. (Castelli, 1935, p. 33.)
Czechoslovakia. The species is still comparatively common in
two well-defined districts. One embraces the mountainous territory
of the Low and the High Tatra, bordered on the west by the Arva
and the Waag Rivers, on the east by the Dunajec and Poprad
Rivers. The other comprises the wooded Carpathians west of the
railway from Munkac to Volovec. According to Dr. Komarec of
ORDER CARNIVORA: CARNIVORES 225
4
Prague, 210 Brown Bears live in this territory, under government
protection. (Tratz, in Castelli, 1935, pp. 8-9.)
Hungary. Blasius (1857, p. 199) reports the bear as still occur-
ring in the Hungary of his time, especially in the Carpathians and
in the Hungarian Erzgebirge. The species is not found in the
reduced Hungary (since World War I) (J. Schenk, in litt., Novem-
ber, 1936).
Yugoslavia. In this country the bear lives especially in the
Gottschee district, Carniola, where it is carefully protected, and in
the immense woods of the Auersperg district, Carniola (Tratz, in
Castelli, 1935, p. 9).
In Croatia about 20 bears are estimated to inhabit the forested
area about Jasenak in the Grosse Kapela. They are also reported
as not rare near Otocac and in the northern Velebit Mountains. The
bears do far less damage than the wolves to livestock, and are
reported as harmless to man. (Wettstein, 1928, p. 33.)
The species occurs in considerable numbers only in Bosnia. It is
found also in Slovenia (forests of Kocevje), in Croatia (forests of
Velebit and Vemika Kapela mountains) , and in some parts of Serbia.
In these regions a total of 272 specimens were killed from 1891 to
1921, including 21 in 1892, 22 in 1893, and 26 in 1910. From 1921
to 1931, 51 specimens were killed. (M. Hirtz, in litt., December,
1936.)
Albania. According to Baldacci (1932-33), the Bear still occurs
commonly in the mountains in the center and north of Albania
(Castelli, 1935, p. 37).
Greece. The bear occurs in Macedonia and Epirus and does not
show a decrease (Game Department, Ministry of Agriculture, Greece,
in litt., October, 1936) . It is not a rarity in the extensive forests of
the Greek and Turkish Balkans (Tratz, in Castelli, 1935, p. 9) .
Bulgaria. "The bears are being killed at all seasons of the year,
in every manner." A 220-kilo specimen was recently killed in a
predatory animal "drive" in the Rhodope Mountains. Reserves
for the preservation of the bear are advocated in the Eastern
Balkans (Stara Planina) and in the Rhodope and the Pirene Moun-
tains. (H. W. Shoemaker, in litt., June 30, 1932, and December
27, 1932.)
The species is found in all the mountains of Bulgaria. The present
number is estimated at about 500. Since 1935 the bear may be
hunted only on a special license, which is issued only for individuals
that have become harmful to cattle pasturing in summer on the
mountains. Protection is assured, and there are some reserves where
hunting is entirely forbidden. (Bulgarian Game Association "Sokol,"
in litt., February, 1937.)
226 EXTINCT AND VANISHING MAMMALS
Rumania. The bear occurs in the coniferous forest zone of the
Banat and the southern Carpathians and in a limited area of the
eastern Carpathians. Toward autumn it comes in search of food to
lower heights, as far as the lower border of the coniferous forests
(Tismana, in the Horjin District, and Brasov). (Calinescu, 1930,
p. 365.)
In the Transylvanian Alps it is still common in some districts
(Tratz, in Castelli, 1935, p. 9) .
Poland. The species was not rare in the eighteenth century, when
it was still found all over Poland. The number is now reduced to
about 250-270 individuals, found chiefly in the eastern Carpathians ;
there are still about 20 in eastern Poland. There are also some in
the Tatra Mountains. The number has perhaps increased of late.
Females and young are absolutely protected, and the hunting of
males is forbidden from January 15 to December 15. (M. Siedlecki,
in Hit., October, 1936.)
About 256 are left in the Carpathians, and 15 in eastern Poland
in the swampy forest of Agarkow (National Council for Nature
Protection, in litt., October, 1936) .
In the future special permits for the shooting of bears will be
granted by the Ministry of Agriculture (Quarterly Information
Bulletin concerning the Protection of Nature in Poland, Kwartal 3,
1935).
The proposed National Parks of the Tatra and of Czarnohora
will be of importance in the protection of bears. The chief aim of
the proposed International Park of Poland, Czechoslovakia, and
Rumania is the creation of a breeding ground for the Bear, Lynx,
Wildcat, Wolf, Stag, etc. (Benedyct Fulinski, MS, 1933.)
Lithuania. The species has been exterminated in this country
since about 1877 (T. Ivanauskas, in litt., November, 1936) .
Latvia. The species is now extinct in Latvia, the last specimens
having been killed in 1880-90 (N. von Transehe, in litt., February,
1937). An occasional straggler comes from Russia or Estonia to
our northeastern forests (Forest Department, Latvia, in litt., March,
1937).
Estonia. About 20 individuals are found in the northeastern
part, in the district of Wirumaa (Wiesland). Hunting is allowed
only on a special permit from the State Forest Department. In
recent years permits have been given for only one specimen each
year. (Zoological Institute, University of Tartu, in litt., October,
1936.)
Russia. In Russia and Siberia the Brown Bear is more or less
generally distributed, and in many areas it has maintained itself
in fairly satisfactory numbers. Under these circumstances it seems
unnecessary here to devote a great deal of space to the local dis-
ORDER CARNIVORA: CARNIVORES 227
tribution. Very detailed information on this point is supplied by
Ognev (1931, pp. 34-108). Various forms have been described from
this vast region; all that are considered valid at all are rated as
subspecies of Ursus arctos by Pocock (1932).
W. G. Heptner writes (in litt., December, 1936) that the species
is found in all forested regions of the U. S. S. R., including Caucasia
and the mountains of Turkestan. In certain regions there are great
numbers. Hunting is allowed in most regions during the whole year,
but in White Russia only on special permit. In one part of Caucasia
and in the mountains of Turkestan hunting is limited to certain
open seasons.
In European Russia, at the present time, the species seems to be
found chiefly in the northern parts, in the Ural region, and in
Caucasia. Many of the records from central Russia seem to date
from the last century, and yet the species still survives near Lenin-
grad and Moscow. In the Caucasus region generally it is quite
common, though rare in Daghestan. (Ognev, 1931, pp. 34-38.) As
many as half a dozen different races have been recognized in Cau-
casia by various authors (Satunin, Smirnov, Lonnberg, Ognev) .
"In the Caucasus, according to Prince Demidoff, it is so common
that the keepers of the Grand Ducal territories have instructions
to treat these animals as vermin, and to kill them whenever occasion
occurs" (Lydekker, 1901, pp. 92-93).
Asia. The Brown Bear is still numerous in many of the thickly
forested areas of Siberia, where the people do not hunt so much now
as formerly. It is distributed from the Urals east through the basins
of the Ob, Yenisei, Lena, and Kolyma to the Anadyr region and
Kamchatka (where it is very common). It ascends to 11,400 feet
in the Sayan Mountains, and to 8,259 feet in the Yablonoi Moun-
tains. (Ognev, 1931, pp. 38-40.) Southward its range extends to
Turkey, Syria, Persia, Afghanistan, the Pamirs, Tian Shan, Hima-
layas, western China, Manchuria, Hokkaido, and the Kuriles. There
is almost a plague of bears in Hokkaido (Inukai, 19326, p. 526).
Many different names (generic, specific, and subspecific) have been
applied to the Brown Bears of various parts of Asia, but Pocock
(1932) regards them as nothing more than races of Ursus arctos.
Separate accounts of two of these forms follow.
The Old World Brown Bear is closely related to the Grizzly Bears
of North America and shows a decided resemblance to them in
food habits and economic status. The considerable human tolerance
exhibited toward it, together with its survival to the present day
in most of the thickly populated countries of Europe, leads one to
question the actual necessity for the ruthless war of extermination
that has been waged upon the Grizzlies in the relatively sparsely
settled areas of the Western United States.
228 EXTINCT AND VANISHING MAMMALS
i
Manchurian Black Bear; Manchurian Grizzly
URSUS ARCTOS LASIOTUS J. E. Gray
Ursus lasiotus J. E. Gray, Ann. Mag. Nat. Hist., ser. 3, vol. 20, p. 301, 1867.
("North China.")
SYNONYM: Melanarctos cavijrons Heude.
FIG.: Sowerby, 1923, pi. 3.
For the purposes of the present report, this subspecies is restricted
to Manchuria and adjacent regions of the Asiatic mainland, although
Pocock (1932, p. 799) provisionally includes with it the bear of Yeso
(Hokkaido) and the Kuriles (U. a. yesoensis Lydekker).
Only five museum skins of this little-known bear seem to be on
record from Manchuria and Mongolia (Pocock, 1932, p. 799). It is
becoming increasingly rare, and calls for government protection in
some way or other if it is to be saved from ultimate extinction
(Arthur de C. Sowerby, in litt., April 24, 1937) .
It is as large as the Kamchatkan Brown Bear (U. a. beringianus
Middendorff) but differs from it on the average, at least, in the
prevalent blackness of its hue. The general color is glossy black;
muzzle brown ; underwool brown. Adult male from Manchuria : head
and body, 6 feet 7 inches; tail, 5.5 inches (Pocock, 1932, pp. 799-
800.)
The range seems to include the forested regions of northern
Manchuria, northern Mongolia, southeastern Siberia, and perhaps
northern Korea.
Sowerby (1920, pp. 230-231) shot a specimen in North Kirin,
Manchuria, and heard reports of a similar animal in South Kirin,
on the lower Sungari River, and in northern Korea. "The specimen
I shot was very savage .... The native Russians and Chinese
greatly fear this animal, as it has been known to kill and devour
hunters."
Sowerby also writes (1923, p. 58) : "The distribution of this
species is doubtful, or, perhaps it would be more correct to say, is
not known. So far it has been recorded only from the forest near
Tsi-tsi-har in South-western Heilung-kiang, and from the forest
in the I-mien-p'o district of North Kirin. From all accounts, how-
ever, it occurs throughout the Manchurian forest, and on into
Primorsk [Siberia]." He adds that a hunter reports this form as
"much rarer than the black bear [Selenarctos] , occurring in the
proportion of one in twenty of the bears shot in the district."
Syrian Bear
URSUS ARCTOS SYRIACUS Hemprich and Ehrenberg
Ursus syriacus Hemprich and Ehrenberg, Symbolae Physicae Mammalium,
decas prima, text to pi. 1, 1828. (Near the village of Bischerre, Mount
Makmel, Lebanon.)
ORDER CARNIVORA: CARNIVORES 229
SYNONYM: Ursus schmitzi Matschie (1917).
FIGS.: Hemprich and Ehrenberg, 1828, pi. 1; Wolf, 1861, pi. 17 (specimen
from the Persian Gulf) ; Pocock, 1937, p. 807, fig.
The Syrian Bear is now extinct, or nearly so, in Palestine and
Lebanon but survives in rather indefinite numbers to the northward
and eastward, where the exact limits of its range have not been
determined.
The type specimen (which was not full-grown) was described as
uniform fulvous-white; it was smaller than Ursus arctos and had
long ears. Its head and body measured 3 feet 8 inches; tail, 6
inches ; height at shoulder, 2 feet 4 inches. Other skins were said to
be fulvous or sometimes almost wholly brown. (Hemprich and
Ehrenberg, 1828.)
The range, according to Flower (1929, p. 149), is "western Asia:
in certain mountainous localities from Asia Minor and Syria to
Persia."
Bodenheimer (1935, p. 114) writes:
The Syrian Bear . . . was not uncommon in N. Palestine in Biblical times.
David boasts of having strangled a bear, which had attacked his herd (I Regum
17, 34) and two bears killed the 42 boys who had scoffed at the prophet
Elisha (II Regum 2, 24). Tristram encountered one in a ravine near Tiberias,
near Beisan and in the Jolan. Schmitz seems to have seen the last specimens on
the southern Hermon (1911, 1913). ... It has not been a menace to flocks of
sheep and goats for a long time, but occasional visits to vine-yards and
fruit-groves are still reported from Syria. The Bear is extinct on the Hermon
and Anti-lebanon, mainly because it was so drastically hunted by German
officers during the war. It is reported to have survived on the Lebanon.
J. C. Phillips writes (in litt., July 20, 1936) that there were
supposed to be a few bears left on Mount Hermon when he was
there in 1912.
The following information, supplied by Dr. William Van Dyck
and Professor West, both of the American University in Beirut, is
transmitted by Theodore Marriner (in litt., 1936) :
"Shortly after the World War, when there were a large number of
army rifles in mountain villages, the number of Syrian bears . . .
was greatly reduced. They were, in fact, exterminated in some parts
of the Anti-Lebanon range, but a few are still reported in the less
accessible parts of both the Anti-Lebanon and Lebanon ranges.
Farther north, in the Gebel Ansariyah and in the Amanus range of
northern Syria and southern Turkey, they are still quite common
in the more wooded sections. At the present time no definite attempt
is being made to preserve the Syrian Bear, although the government
policy of forbidding civilians to carry rifles indirectly helps towards
this end."
Aharoni (1930, pp. 336-337) gives the following account (some-
what freely translated) : "During the war, while stationed in Leba-
230 EXTINCT AND VANISHING MAMMALS
non, I found that the light Isabella-colored bear [Ursus syriacus
H. and E.] , with the dimensions of the original description, inhabited
only the green shrubbery of the Anti-Lebanon, while the smaller
brown bear [U. syriacus schmitzi Matschie] inhabited only the bare
snow-fields of the Lebanon. I saw examples of both subspecies in
nature and still have specimens from Lebanon. To-day the bear
has disappeared not only from Palestine, but perhaps also from
Syria.
"Last year I became convinced that the Mesopotamian bear in-
habiting the Jebel Abdul-Aziz [in the present Syria; lat. 36 30'
N., long. 40 30' E.] represents a distinct subspecies."
F. S. Bodenheimer writes (in Hit., March, 1937) that the animal
is now extinct in Palestine and Lebanon but probably still survives
in Anti-Lebanon in small numbers. He adds that protection is most
highly desirable.
Pocock states (1932, p. 793) that "the bears of Asia Minor and
Syria merely differ from the typical Brown Bear of Europe in being
on the average paler in colour, intermediate specimens occurring
in the Caucasus and perhaps in northern Persia." He records speci-
mens from Smyrna and from Sumela, 30 miles south of Trebizond,
Turkey.
Blanford (1876, pp. 46-47) gives the following account of bears
in Persia:
"Major St. John, . . . who has seen several Elburz bears, assures
me that, although they are darker than the true Ursus Syriacus
which is found in Southern Persia, they are much paler in colour
than the common bear of Europe. . . . '
Ursus syriacus "is, as Major St. John assures me, the bear of
South-western Persia. It is not the bear of Baluchistan, but is said
to be found between Bampur and Bam. It is found* pretty commonly
in the neighbourhood of Shiraz and in the hills bordering on Meso-
potamia."
To this St. John adds (in Blanford, 1876, p. 47) : "This bear is
found throughout the mountains of Western and Northern Persia,
possibly extending to Khorassan. In many places watchers are set
at night to keep the bears from the ripening grapes."
Atlas Bear; Crowther's Bear
URSUS CROWTHERI Schinz
Urs[us~\ Crowtheri Schinz, Synopsis Mammalium, vol. 1, p. 302, 1844. (Based
upon "the Bear of Mount Atlas," Blyth, Proc. Zool. Soc. London 1841,
p. 65; type locality, "the foot of the Tetuan mountains, about twenty-five
miles from that of the Atlas.")
The bear of North Africa is almost a mythical species, for no
specimen has ever reached a museum. No very definite news of the
ORDER CARNIVORA: CARNIVORES 231
species has been obtained for nearly a century, and it is very prob-
ably extinct.
An adult female was smaller than the American Black Bear but
more robustly formed and with a shorter and broader face, though
the muzzle was pointed; toes and claws remarkably short; hair
black or brownish black, shaggy, 4 or 5 inches long; muzzle black;
under parts orange-rufous (Blyth, 1841, p. 65) .
Pliny, though skeptical himself, quoted Roman annals to the
effect that Domitius Ahenobarbus, an aedile of 61 B. C., had shown
in the Roman arena a hundred Numidian bears, conducted by as
many Ethiopian hunters. The bear of Libya was mentioned by
Pliny's contemporaries, Juvenal and Martial, and a long time pre-
viously by Virgil. (G. Cuvier, 1825, vol. 4, pp. 325-326.)
Strabo says expressly that the Moors dressed themselves in bear
and lion skins (Wagner, 1841, p. 70).
Blyth (1841, p. 65) gives a brief description of the animal, based
upon information supplied by Mr. Crowther, who had spent some
time in Morocco. "Upon questioning Mr. Crowther respecting the
Bear of Mount Atlas, which has been suspected to be the Syriacus,
he knew it well, and it proves to be a very different animal. . . .
This individual was killed at the foot of the Tetuan mountains,
about twenty-five miles from that of the Atlas. It is considered a
rare species in that part, and feeds on roots, acorns, and fruits. Does
not climb with facility; and is stated to be very different-looking
from any other Bear." An unsuccessful effort was made to preserve
the skin of the specimen mentioned.
According to Loche (1867, p. 52), Shaw (1743) mentions a bear
in the Atlas Mountains. Loche also states that the Emperor of
Morocco had recently sent to the zoological garden of Marseilles a
live bear coming from his territory.
Bourguignat (1867, pp. 41-46) contributes the following informa-
tion. Herodotus records a bear from western Libya. Poiret, a French
botanist and zoologist, reports (1789) bears from the Atlas Moun-
tains, and mentions a fresh skin brought by an Arab into Mazoule.
A friend of Bourguignat's, M. Letourneux, had reports of many
bears in the region of fidough, and learned of others occurring not
long previously on Djebel-Bou-Abed, Djebel Gherar, Djebel Debhar,
and Djebel Thaya, Algeria. The animal was said to be small, thick-
set, and brown, with a white spot on the throat, and to be very fond
of honey and fruits. Bourguignat himself records skeletal remains
of a bear from a cavern on Djebel Thaya in the Province of Con-
stantine, to which he gives the name of Ursus jaidherbianus. Human
artifacts associated with these remains were believed to date from
the early Christian Era.
232 EXTINCT AND VANISHING MAMMALS
Lataste (1885, pp. 235-237), in reviewing the evidence for the
presence of bears in Barbary, considers that the case has by no
means been proved.
"Since Mr. Crowther's time no more definite news has been re-
ceived of this bear, though other travellers have reported statements
of Arabs and Moors that such a creature exists in the mountains
of Eastern Morocco and Western Algeria" (Johnston, in Bryden,
1899, p. 608).
"In view of the apparent rarity of the animal, it is important
to mention that fossilised remains of bears have been discovered in
caverns in north-western Africa, as well as in the rock-fissures of
Gibraltar" (Lydekker, 1908, p. 463).
During the years 1892-96, "fchere were still rumours of Bears
(Ursus crowtheri) in the Western Atlas, but although they certainly
existed there in the first half of the last century I have never heard
of one being killed or seen since this region became better known
with the penetration of the French into Morocco, though there may
be a possibility that a few exist" (Pease, 1937, p. 81).
The foregoing accounts seem to constitute fairly strong evidence
of the former existence of a bear in North Africa. It must be
acknowledged, however, that no less an authority than Cabrera
(1932, pp. 10, 102-103) throws the whole case for the Atlas Bear
out of court. But he is hardly correct in maintaining that its sole
basis is the "fantastic" account of Blyth.
On first thought, the Atlas Bear might appear to be a note-
worthy exception to the general rule that recently extinct mammals
have succumbed to the advance of the European type of civilization.
Yet one of the tools of that civilization, the rifle, in the hands of
the Moors, must have at least contributed to the animal's downfall.
Nevertheless, the disappearance of the Barbary Lion from Morocco
in the early part of the present century is singular enough (Cabrera,
1932, p. 186) , and the still earlier disappearance of the Atlas Bear
is even more puzzling.
Family MUSTELIDAE: Weasels, etc.
This family is distinguished by the large number of valuable fur-
bearers represented in it; and many of the species have been seri-
ously reduced by the demands of the fur trade. Its distribution is
practically as cosmopolitan as that of the Canidae; it extends to
Borneo and the Philippines, but not to Australia. There are about
35 genera and 400 species and subspecies. Nineteen forms (including
one extinct species) are discussed in Dr. Allen's volume on the New
World (1942), and nine forms of the Old World in this volume.
ORDER CARNIVORA: CARNIVORES 233
European Mink. Vison (Fr.). Norz; Sumpf otter (Ger.)
MUSTELA LUTREOLA Linnaeus
[Mustela] Lutreola Linnaeus, Syst. Nat., ed. 12, vol. 1, p. 66, 1766. (Finland.)
FIGS.: Royal Nat. Hist., vol. 2, p. 68, 1894; Martin, 1910, pi. 33; Didier and
Rode, 1935, p. 303, fig. 176.
The European Mink is a rare and vanishing species in central
Europe and France but is more generally and more commonly dis-
tributed in Russia.
The general color is a rich dark brown; region about the mouth
whitish; tip of the tail blackish. Head and body, 350 (female) to
400 mm. (male); tail, 130 (female) to 140 mm. (male). (Miller,
1912, pp. 415, 418.)
The range extends from western France eastward to the Tobol
and Irtish Rivers in western Siberia; south to Austria, Hungary,
Rumania, and Transcaucasia ; north to Finland and northern Russia.
France. Though recorded by Lesson in 1840 in Poitou and Sain-
tonge, the Mink was long overlooked in France. It seems to have
been formerly rather common in the center, the west, the southwest,
Normandy, and the Vosges. The present range consists of a narrow
zone extending from the Jura to the vicinity of Nantes and in a
general way following the valley of the Loire. Here the species
seems to become rarer and rarer. The decrease is due to the active
hunting of the animal, for its fur is very valuable and it is also
considered a harmful species. Prohibition of hunting and surveil-
lance of the fur trade would be the only means of conservation;
but these measures would be very difficult to apply. (E. Bourdelle,
in litt., March 6, 1937.)
Martin (1910, p. 35a) extended the range to the Gironde and to
Brittany.
The Mink's food includes fish, frogs, crawfish, ducks, and small
mammals (Didier and Rode, 1935, p. 304) .
Germany. The species has been exterminated in western Ger-
many and is very rare in northern and eastern Germany. Latest
dates of capture are: Mecklenburg, 1894-96; Hannover, 1902; East
Prussia, 1909; Liineburger Heide, 1910. At present there is no open
season. (Internationale Gesellschaft zur Erhaltung des Wisents, in
litt., October, 1936.)
It had disappeared from Schleswig-Holstein by about 1890 (Mohr,
1931, p. 32).
During recent years solitary individuals are still regularly shot or
seen in the east (Krumbiegel, 1930, p. 6) .
Switzerland. Fatio (1869, p. 336) has only a few doubtful records
from this country.
Austria. The Mink was formerly found in Burgenland and prob-
234
EXTINCT AND VANISHING MAMMALS
ably also in Lower Austria. It is now absolutely protected in Bur-
genland, where it is said to still exist, though reliable reports are
not obtainable. (G. Schlesinger, in litt., March, 1937.)
Czechoslovakia. The known specimens are mostly from the
Carpathians, in the former Hungary (J. Schenk, in litt., November,
1936).
Hungary. From the present limits of Hungary (since the World
War) only one specimen is known. The animal has no legal pro-
tection. (J. Schenk, in litt., November, 1936.)
FIG. 23. European Mink (Mustela lutreola). After Lydekker.
Rumania. The Mink is common, like the Otter, but has a greater
distributional area, since it is more adaptable to civilization (Cali-
nescu, 1930, p. 366) .
Poland. It was formerly quite common all over Poland but is
now very rare, being found especially in the southeast, in Polish
Podolia and in the Eastern Carpathians. Hunting is forbidden
from February 1 to December 31. Lately there has been a demand
for complete protection. (M. Siedlecki, in litt., October, 1936.)
Kuntze (1935, p. 63) records it from northeastern and south-
eastern Poland.
Lithuania. The species is exterminated except in the eastern
part of the country, where it is still found in the districts of Zarasai
and Utona. The annual production amounts to as many as 150
skins. The value of one is about 40 Litas (5-6/) . So far no protec-
tive measures have been adopted. (T. Ivanauskas, in litt., Novem-
ber, 1936.)
Latvia. In 1908-09 it was reported as numerous in Courland
and widely distributed in Livonia (Ognev, 1931, p. 759).
ORDER CARNIVORA: CARNIVORES 235
The present stock is about 2,000 individuals. The increase through
natural propagation is not important, and the annual kill is about
300-400 specimens. The species is protected from March 1 to Novem-
ber 14. Forest guards are not allowed to kill it. (Forest Department,
Latvia, in litt., March, 1937.)
Estonia. The species is found throughout the country in suitable
areas. There is a steady decrease, owing to the drying up of the
country. The animal is not threatened by man, but it is without
any legal protection. (Zoological Institute, University of Tartu,
in litt., October, 1936.)
Finland. The species is apparently distributed in the southern
part of the country (Ognev, 1931, p. 758).
Russia. From Ognev's data (1931, pp. 758-761), the Mink ap-
pears to be widely and more or less commonly distributed over most
of Russia, from Kandalaksha Bay, the lower Dvina, and the Pet-
chora and Usa Rivers in the north to the Ukraine, the Caucasus,
and Astrakhan in the south. Westward it is found about Lake Onega,
in Volhynia and Podolia, and on the lower Dniester. At the bazaar
of White Russia 473 Mink skins were sold in 1926-27, and 649 in
1927-28. Eastward the species is found in the Ural region, from
the tributaries of the Petchora in the north to Orenburg and the
Ilek River in the south. Beyond the Urals it extends only to the
Tobol and Irtish Rivers. In Transcaucasia it occurs on the Bzyb
River.
The animal is strongly persecuted as a fur animal and is rare
in certain regions. Hunting is not allowed in the Volga region and
in the eastern part of European Russia. There is no danger of ex-
tinction, except in certain industrial regions. (W. G. Heptner, in
litt., December, 1936.)
Russian Sable. Marte zibelline (Fr.). Zobel (Ger.)
MARTES ZIBELLINA (Linnaeus)
Mustela zibellina Linnaeus, Syst. Nat., ed. 10, vol. 1, p. 46, 1758. (Northern
Asia; type locality restricted by Ognev (1925, p. 276) to "the northern
part of the government of Tobolsk.")
FIGS.: Royal Nat. Hist., vol. 2, p. 55, 1894; Ognev, 1931, pi. 5 (M. z.
sahalinensis) ; Zeitschr. f. Saugetierk., vol. 9, pi. 18, fig. 7, 1934 (Amur
form) .
This is one of the animals that has suffered particularly from
the "curse of beauty." It has been decimated by the demands of
the fur trade and has disappeared from considerable areas within
its former range. Its principal home is in Siberia.
The Sable bears considerable resemblance to the Pine Marten
(Martes martes). It has a cone-shaped head, large ears, a bushy
f
236 EXTINCT AND VANISHING MAMMALS
tail, and comparatively stout limbs. The fur is thick and soft; the
color varies from blackish, mixed with gray and brown, to yellowish
brown; throat sometimes orange. Head and body, about 20 inches;
tail, about 7 inches.
The former range of the species included the forested regions
from northern Russia east to the Anadyr district, Siberia; it ex-
tended south to the southern Urals, the Altai and Sayan Mountains,
Manchuria, the Ussuri district, Hokkaido, Sakhalin, and Kamchatka.
The following subspecies have been recognized (cj. Ognev, 1925,
and 1931, pp. 560-598) , but the ranges so far assigned to them do
not cover the entire range of the species:
Martes zibellina zibellina (Linnaeus). (Original reference and
type locality given above.) Tobolsk Sable.
Color dull and pale, varying from cinnamon-drab to pale brownish
yellow and even to dark brown; underfur light and dull.
Range: the Ob Basin and the Ural region.
M. z. yeniseensis Ognev, Jour. Mammalogy, vol. 6, p. 277, pi. 26,
fig. 3, 1925. ("Krasnoyarsk district, the forest on the plain along
the Yenisei River," Siberia.) Yenisei Sable.
Color more dusky warm brown in comparison with the Tobolsk
Sable.
Range: the great forests of the districts of Krasnojarsk, Ashinsk,
and Kansk, in the Yenisei Basin.
M. z. sajanensis Ognev, Jour. Mammalogy, vol. 6, p. 278, 1925.
("Orsyba River, northern part of the Sajansky Mountains," Siberia.)
Sayan Sable.
General color dark brown; underfur pale yellowish.
Range: "the mountain country of the rivers Uda, Kasyr and
especially of the Kasyr-Suk and partly of the Usa."
M . z. princeps (Birula) . (Mustela zibellina princeps Birula, Ann.
Mus. Zool. Acad. Imper. Sci. Petrograd, vol. 22, p. 08, 1922; the
mountain country of Bargusin, Transbaikalia, Siberia.) Bargusin
Sable.
Fur soft and silky; color a brilliant blackish brown; underfur
bluish gray, brownish at bases and tips; throat patch much reduced,
commonly not visible.
Range: mountain forests, Bargusin Hills and spurs of the Stano-
voi Mountains, Transbaikalia.
M. z. kamtschadalica (Birula). (Mustela zibellina subsp. kamt-
schadalica Birula, C. R. Mus. Zool. Acad. Sci. Petrograd 1918,
p. 82 (fide Ognev) ; Kamchatka.) Kamchatka Sable.
General color between warm sepia and mars brown; underfur
pale yellowish gray ; skull large.
Range: Kamchatka.
M. z. sahalinensis Ognev, Jour. Mammalogy, vol. 6, p. 279, pi. 26,
fig. 4, 1925. ("Saghalien, Wedernikovo.") Sakhalin Sable.
ORDER CARNIVORA: CARNIVORES 237
Winter pelage like that of the Kamchatka Sable but of a more
decided cinnamon tint and lighter; throat patch of the same cinna-
mon color; head avellaneous, back darker; flanks sayal brown or
tawny-olive; underfur pale yellowish, more cinnamon at the tips;
summer pelage duller and darker, more brownish.
Range: the whole of Sakhalin Island.
M. z. brachyura (Temminck). (Mustela brachyura Temminck,
in Siebold, Fauna Japonica, Mammiferes, p. 33, 1844; Matimaja,
Hokkaido, Japan.) Japanese Sable.
FIG. 24. Russian Sable (Maries zibellina subsp.)
Inferior to the Siberian Sable in fineness and length of fur; back
and tail dark brown; sides and limbs lighter; long hair of feet con-
cealing the claws. Tail, 3.5 inches. (Temminck, 1844, pp. 33-34.)
Range : Hokkaido and the Kuriles.
Russia. In past centuries the Sable's range extended westward
perhaps as far as the Kola Peninsula or even Lapland. In the six-
teenth and seventeenth centuries it was found on the Petchora
River and probably at the same time in the Dvina region. In Pallas's
time it occurred in the vicinity of Ufa, west of the southern Urals;
the last one in this general region was killed in 1850 near Ufimsk.
Its southern limit on both slopes of the Urals was about latitude
52 N., or possibly 51 N. About 1700 it inhabited the entire Gov-
ernment of Perm and the eastern half of the Governments of Vo-
logda, Archangel, and possibly Viatka. By 1875 about 300 Sables
were trapped annually in the northeastern part of Perm. More
238 EXTINCT AND VANISHING MAMMALS
recently the Russian population of the species seems to have become
largely restricted to the Urals, where it is less numerous on the
western slopes than on the eastern. However, in 1925 it was still
common on the Ilych River, a tributary of the Fetch ora. (Ognev,
1931, pp. 569-570.)
The Ural form is considered distinct from M. z. zibellina but is
not named. It occurs sporadically and rarely along the western
foothills (headwaters of the Shugora, Laga, Ilych, and Petchora) ;
it is also rare on the eastern slopes, at the sources of the Losva,
Aspia, Purma, Ushma, Toshemok, and Wishaj Rivers. The fur is
considered the finest in western Siberia. (Ognev, 1925, p. 277.)
Siberia, Ob Basin (M. z. zibellina). In the Government of To-
bolsk the Sable is not rare in the taiga forests of the Pelym River;
it is rare along the Tavda River and in the Tarsk and Surgut dis-
tricts. At the end of the last century more than 300 Sables were
obtained annually along the Jugan River. The species is absent
between Beresof and Obdorsk. Along the Rivers Omi, Tara, and
Irtish, and in the southern part of the Government of Tobolsk it
was very scarce in 1886. In the Narym district it was numerous on
the upper Wasugan River in 1875. It occurs on the Ket and Chulym
Rivers, and was particularly numerous on the Tchirk-Ul River about
1923. It avoids the steppes in the central part of the Government of
Tomsk. (Ognev, 1925, p. 277, and 1931, pp. 571-572.)
Prejevalsky (1879, p. 233) reported the Upper Katuna, the Bukh-
tarma, and their tributaries, in the Russian Altai, as particularly
good districts for Sables. The hunters used specially trained dogs,
and endeavored to surround the animals with nets, which were as
much as 1,000 feet long and 4 feet high. The average price of a
sable skin was then 15 rubles.
Siberia, Yenisei Basin (M. z. yeniseensis, M. z. sajanensis, M. z.
princeps) . The Sable is found in suitable areas from the Mongolian
boundary northward to latitude 69 N. The form living along the
Tunguska River and near Turukhansk probably represents an un-
described subspecies ; the same form is found in small numbers in the
adjacent Khatanga Basin. The species is less common in the Gov-
ernment of Irkutsk than in the Government of Yenisei. It is absent
from the steppes in the vicinity of Minusinsk, Achinsk, and Kras-
noyarsk. It is common in the Sayan region on the Kasyr-Suk and
Uda Rivers, and occurs on practically all sides of Lake Baikal.
According to Turov (1923) , 700 skins were exported annually across
Bargusin from the Verkhne Angarsk and Podlemorsk districts near
Lake Baikal. (Ognev, 1931, pp. 572-573.)
Siberia, Lena Basin. In the Olekma-Vitim mountainous country
the Sable is very rare. Far to the north, in the enormous region
between the Anabar, the Olenek, the Lower Tunguska, and the Vilui,
ORDER CARNIVORA: CARNIVORES 239
it disappeared about the middle of the last century. About 1873
it was found occasionally on the Patom River. Its disappearance
from the entire southern Muisk district was unusually rapid. It
has been recorded from the Aldan and Mae Rivers. (Ognev, 1931,
p. 573.)
The dark animals furnish the most valuable skins; in peace times
they fetched as much as 2000 marks. They come mostly from the
Vitim Plateau and from the Bargusin district, where the Sable is
now almost exterminated. (Klemm, 1930, p. 367.)
Eastern Siberia. At present the Sable is not found in the Verk-
hoyansk and Kolyma districts. In former times it was widely
distributed along the Kolyma and Omolon Rivers, but it finally
disappeared from the Kolyma district in 1852. The species has
long since vanished from the Anadyr River region ; the last one was
found near the village of Eropol about 1847.
By 1900 the species was rare in the Gizhiginsk district, though
in former years from 30 to 50 Sables were collected annually, par-
ticularly from the Penzhina Valley and from northern Kamchatka.
Possibly this form belongs to M. z. kamtschadalica, which is widely
distributed in Kamchatka, especially in the Petropavlovsk district.
(Ognev, 1931, pp. 574-575, 595.)
In Kamchatka the Sable was decimated in Dybowski's time (1879-
85). At the beginning of the nineteenth century a hunter could
get 40 animals a day, and the annual production of Kamchatka
amounted to 10,000 skins. The natives did not endanger the stand
of Sables, but by 1881 Cossack and Tungus immigrants reduced the
yield to 2,883. The abundance of the animals in some years was
dependent on the wholesale occurrence of a vole, Microtus oecono-
micus. (Kuntze, 1932, p. 47.)
In the western Amur region the species occurs on the Argun and
Shilka Rivers. In the middle and lower Amur Basin, the Sables
from the Albazin area, the Zeya River, and the Bureya Mountains,
which are very dark in color, and costly, may belong to the sub-
species M. z. princeps. In 1861 the species was reported as particu-
larly numerous on the Amgun River. In the Ussuri district it varies
from common to rare, and has even disappeared entirely in some
parts. The Ussuri Sable is probably very near to M . z. sahalinensis.
(Ognev, 1925, pp. 279-280, and 1931, pp. 573-574.)
Sowerby (1923, pp. 63-65) says that among the Tartars of the
Primorsk coast in southeastern Siberia, "sable hunting is their chief
end and aim in existence." He continues:
It is certain that it was largely the presence of the sable throughout Siberia
and in the Amur and Primorsk that led the Russian pioneers and conquerors
across that wide stretch of country. . . . Thus we must look upon this little
animal as having a very important bearing upon the history of these regions.
240 EXTINCT AND VANISHING MAMMALS
. . . The only trouble is that with the unrestricted hunting that takes place, and
the steady increase in the settlement of the country, this valuable supply of
fur-bearing animals is rapidly diminishing. . . .
It is evident that one of the chief objects of the conquest of Siberia was
to secure a supply of sable skins for the Imperial Government, and it is
significant that the conquering Cossacks . . . always imposed a heavy tribute
of sable skins upon the Tartar tribes they defeated, and brought under
subjection ....
In Siberia this animal is protected by the Government, and comparatively
recently it was given a five years closed season.
The Cedar Valley Reservation (Kedrovaya Pad) on Amur Bay,
comprising 7,500 hectares, and the Kronotsk Bay Reservation in
Kamchatka, comprising 15,000 hectares, provide for the protection
of the Sable (Makaroff, in Skottsberg, 1934, pp. 433-434) .
In Sakhalin the Sable is distributed over the entire island, and in
1889 it was considered more numerous there than in any other part
of Siberia (Ognev, 1931, p. 574).
Mongolia. P. P. Sushkin reported in 1925 that the Sable was a
regular inhabitant of the southern slopes of the Altai Mountains,
about the headwaters of the Black Irtish and the Urungu. It is also
abundant in the vicinity of Kossogol, at the southern base of the
Sayan Mountains. (Ognev, 1931, p. 572.)
Manchuria. The most valuable fur-bearing animal of Manchuria
is the Sable (Sowerby, 1934, p. 286) .
"The Manchurian sable does not come up to those from the Amur,
Primorskaya and Siberia in the value of its pelt. ... It is said
that the Chinese nearly always hunt the sable by running it down
with dogs. . . .
"Unfortunately the Chinese Government is not alive to the value
of its game and fur-bearing animals and birds, and so affords no
manner of protection. There can be only one result of this; com-
plete extinction of the sable in the provinces of Heilungkiang and
Kirin." (Sowerby, 1923, pp. 64-65.)
Japan. Temminck (1844, pp. 33-34) described the Japanese
Sable from Yezo (Hokkaido) and added that it was common in all
the Kuriles. It was being utilized in the fur trade in his day.
In Hokkaido "the sables . . . have . . . met a sad fate and in
spite of particular protection, their coming back to their existence
as before is anything but promising. . . .
"It is interesting to note that the increase of the Japanese minks
[Mustela itatsi] in Hokkaido associates closely with the decrease
of the sables which occupied the land before the minks and de-
creased inland gradually from the southern part. The number of
sables caught in Hokkaido was 2,395 in 1906, 765 in 1910, 1,706 in
1915 and 214 in 1919 respectively." (Inukai, 19326, pp. 524, 527.)
Uchida reports (1935, p. 8) a total of 5,948 Japanese Sables taken
during the six-months open season of 1929-30.
ORDER CARNIVORA: CARNIVORES 241
In the Kuriles the Sable is threatened with destruction (Miyoshi.
in Skottsberg, 1934, p. 412) .
Economics and conservation. The Sable has been very actively
hunted for several centuries. Its numbers are now much reduced,
and in only a few regions can it be said to be "not rare." Its range
also has been considerably reduced and has become discontinuous.
There has been shrinkage of the range at its western limits in Russia
and at its southern limits in western and southern Siberia. From
time to time hunting has been forbidden in the whole or in certain
parts of its range. These measures have given good results, and the
decrease in numbers has been halted for several years. In order to
obtain an increase, a closed season has been maintained on the
whole territory of the U. S. S. R. Several great reserves have been
created to afford protection to the Sable (Barguzinsk, Kronotski,
Sikhote-Alin, Kondo-Sosva) . Successful propagation has been car-
ried out, and several "sovkhoz" have been specially created for
sable-farming. (W. G. Heptner, in litt., 1937.)
In 1928 the entire Sable production of Asia was 15,000. In 1929-30
the yield in that part of eastern Siberia bordering Manchuria was
1,925 skins (Kuntze, 1932, p. 47).
Wolverine; Glutton. Glouton arctique (Fr.) Vielfrass (Ger.)
GULO GULO (Linnaeus)
[Mustela] gulo Linnaeus, Syst. Nat., ed. 10, vol. 1, p. 45, 1758. (Lapland.)
FIGS.: Fitzinger, Bild.-Atlas, Saugth., fig. 70, 1860; Royal Nat. Hist., vol. 2,
p. 71, fig., 1894; Zeitschr. fur Saugetierk., vol. 8, pi. 31, 1933; Oguev,
1935, pi. 2.
The range as well as the numbers of the Wolverine have become
reduced in northern Europe, but its status in northern Asia has
remained more satisfactory.
The general form is heavy and badgerlike; fur long and dense;
tail bushy; general color a rich dark brown, becoming blackish on
legs, feet, and tail ; a broad light brownish or yellowish band across
rump and upper side of basal part of tail, extending forward to
shoulders, where it gradually disappears; a cream-buff area across
the head between ears and eyes. Head and body, 825 mm.; tail,
125 mm. (pencil, 75). (Chiefly from Miller, 1912, pp. 434-440.)
"In prehistoric times, the wolverine was found in England, and
indeed ranged as far south as the Pyrenees" (Lydekker, 1901,
p. 112).
Its recent range is given by Miller (1912, p. 434) as the "northern
forests of the Old World; in Europe, confined to Scandinavia and
northern Russia." It is stated by Trouessart (1910, p. 71) a little
more fully: "Circumpolar Europe (but not the islands north of the
9
242 EXTINCT AND VANISHING MAMMALS
continent), south to lat. 55 N. (Lithuania, Volhynia, and northern
Germany where it is now exterminated) ; in Asia south to the Altai.
Still lives in northern Norway, Sweden, and Lapland."
Norway. In former times it was common throughout the country,
but only in the mountains in the southern part. At present it is
rare in the high mountains and probably will soon become com-
pletely extinct. Large bounties are paid for every specimen because
of the damage the Wolverine does to cattle and reindeer. (Hj . Broch,
in litt., December, 1936.)
It is supposed that a small stock is left in the south of Norway
on Hardangervidda and in Jotunheimen. While still found in the
northern parts, it shows a considerable decrease there. It is ques-
tionable whether it is possible to preserve the Wolverine in Norway.
(Director of Forestry, Ministry of Agriculture, Norway, in litt.,
January, 1937.)
Sweden. The Wolverine inhabits chiefly the mountain forests
and the areas above the tree limit. It has therefore been mostly
restricted to the country northward from northern Dalecarlia (lat.
61 30' N.). In Wermland it was formerly found in small numbers,
but disappeared before the middle of the last century. Single speci-
mens have even been found as far south as Scania. At present, it
occurs only from Jemtland northward.
The skin of the Wolverine has been valued for centuries, and the
animal has also been pursued because of its damage to livestock,
mostly reindeer but also sheep to some extent. Consequently boun-
ties are paid, and have recently been increased; the State pays
10 Cr., while the Lappfund pays 100 Cr. for old animals and 50 Cr.
for cubs. The total number of Wolverines killed in the whole
country is, by decades, as follows:
1856-1865 1,159 1896-1905 1,084
1866-1875 1,201 1906-1915 717
1876-1885 1,240 1916-1925 639
1886-1895 992 1926-1934 517
There is a pretty steady decrease in numbers from the beginning
of the present century, indicating that there is danger of extermina-
tion. (Einar Lonnberg, in litt., October, 1936.)
Finland. Ognev (1935, p. 95) mentions the occurrence of the
Wolverine about Lake Enara.
Latvia. In the Baltic states the species was once found in large
numbers, but now seems to have disappeared. In 1875 a specimen
was killed near Gerki in Courland, and in 1876 another near Jacob-
stadt. (Ognev, 1935, p. 94.)
Lithuania. Its former existence is uncertain (T. Ivanauskas,
in litt., November, 1936) .
ORDER CARNIVORA: CARNIVORES
243
Poland. It was quite frequent in the eighteenth century and even
in the nineteenth century, but is now most probably exterminated
(M. Siedlecki, in litt., October, 1936). It lived till the end of the
last century in Volhynia, Podolia, and Polesia (Lubicz v. Nieza-
bitowski, 1934, p. 190) .
Germany. Two reports of Wolverines in central or northern
Germany in the eighteenth century are evidently based upon escaped
captives (Blasius, 1857, p. 211; Hilzheimer, 1933, pp. 219-221).
Russia. The species formerly ranged southward to the northern
Ukraine. At present it is found rarely in the Western Area, and pos-
FIG. 25. Wolverine (Gulo gulo)
sibly in Volhynia. Once thought to have been exterminated in White
Russia at the end of the nineteenth century, it may have survived
till later. It formerly occurred in the Governments of Novgorod
and Olonets. It ranges northward to the Kola Peninsula. Prior
to 1901, 200-300 Wolverines were collected annually in the Govern-
ment of Archangel, but at present there is a decrease. Prior to 1925,
75 animals were captured annually in the Petchora district. The
species now seems to be very rare in central Russia (Ivanovo Indus-
trial Area and adjacent areas) . It is widely distributed in the Urals,
south to about lat. 53 N. (Ognev, 1935, pp. 94-96.)
Siberia. The range of the Wolverine extends from the Urals
east to the Anadyr district and Kamchatka ; on the north it reaches
the Arctic coast in places; on the south it extends to the Altai Moun-
tains, the Tannu-Ola Mountains (Mongolia) , the Bargusin district,
the Amur region, the Ussuri district (lat. 44 N.), and Sakhalin.
Over this vast area its status varies considerably; it is reported as
numerous in some places and as rare or absent in others: (Ognev,
1935, pp. 97-100; map, p. 101.)
244 EXTINCT AND VANISHING MAMMALS
The Wolverine is widely distributed in the forested regions of
the U. S. S. R. as a whole (Russia and Siberia) . It is less common in
northern Russia than in Siberia. Its decrease in central Russia
results from the decrease of the forest cover. It is very common in
parts of Siberia. It causes great damage to hunting interests, taking
animals caught in traps, destroying hunters' provisions, attacking
young ungulates and even adults in deep snow. It is not legally
protected and may be killed at any time. (W. G. Heptner, in litt.,
December, 1936.)
Sakhalin. "Schrenck and Kishida reported it from the island. It
is ... a rare 'animal on Sakhalin." (Kuroda, 1928, p. 227.) A speci-
men was taken in 1934 (Kuroda, 1938, p. 26) .
Manchuria. "I heard sufficient from authentic sources to con-
vince me that the animal is fairly common, at least in the northern
forested area" (Sowerby, 1923, p. 71).
Mongolia. The species is reported in the Tannu-Ola Mountains
(Ognev, 1935, pp. 98, 100) . "I once saw a skin from the Urga district
in Northern Mongolia" (Sowerby, 1923, p. 71).
Economics. "Wolverine fur has been much in vogue of late
years, and has consequently appreciated in value. For a good skin,
thirty shillings is often asked." (Lydekker, 1901, p. 112.)
The world's fur production for 1928 included 6,000 Wolverine
skins [some probably from North America] (Jour. Soc. Preservation
Fauna Empire, pt. 12, p. 64, 1930) .
Folklore. Many curious bits of folklore concerning the Wolverine
have been handed down. Some of them are quoted by Lloyd (1854,
pp. 16-18) from Pontoppidan and Olaus Magnus.
Family VIVERRIDAE: Civets, Mongooses, etc.
This Old World family ranges over southern Europe, Africa,
Madagascar, southern Asia, and the Malay Archipelago as far as
Timor, Ceram, and the Philippines. There are about 40 genera and
350-400 forms. Accounts of six forms are given here.
Malay Binturong; Bear-cat; Black Marten. Bintoeroeng
(Dutch)
ARCTICTIS BINTURONG BINTURONG (Raffles)
Viverra? Binturong Raffles, Trans. Linnean Soc. London, vol. 13, pt. 1, p. 253,
1821. ("Malacca,")
FIGS.: Geoffroy and Cuvier, Hist. Nat. Mammiferes, vol. 5, pis. 201, 202
(subsp.?), 1824; Sclater and Sclater, 1899, p. 128, fig. 26; Lydekker, 1900,
pi. 9, fig. 1 (subsp.?).
Opinions differ as to the rarity of the Binturong, the six subspecies
of which range from northeastern India and Tonkin through the
ORDER CARNIVORA: CARNIVORES 245
Malay Peninsula to Sumatra, Java, Borneo, and Palawan. Never-
theless, it is evidently in need of total protection, more especially
because of the demands upon it by the Chinese for medicinal pur-
poses.
There has long been uncertainty as to the exact taxonomic status
of the various described forms, owing chiefly to the inadequacy of
the series of museum specimens. The species as a whole is dis-
tinguished from all other Viverridae by its long ear tufts and by
the prehensile nature of its long, bushy tail. The fur is long, coarse,
and black, more or less washed with gray, fulvous, or buff. (Lydek-
ker, 1893-1894, p. 463.) The present subspecies is larger and darker
than A. b. penicillatus; its winter coat is much shorter and less
luxuriant with underwool than that of albijrons, and the long hairs
are less extensively annulated with lighter color (Pocock, 1933,
p. 1030). Body, 30 inches; tail, nearly the same (Raffles, 1821,
p. 253).
The Malay Binturong inhabits the Malay Peninsula north to
Tenasserim and Siam, and also Sumatra.
Siam. In this country the Binturong seems to be rare (Gylden-
stolpe, 1919, p. 148) . Specimens have been recorded from Sikawtur,
northwest of Raheng, western Siam; from Prachin, central Siam;
from Sai Yoke, southwestern Siam; and from Bang Nara, Patani,
Peninsular Siam (Kloss, 1917, p. 293, and 1919, p. 53; Gyldenstolpe,
1919, p. 148). In Ratburi Province, southwestern Siam, "the
Karangs are well acquainted with the animal and state it is
generally distributed in evergreen forest" (Gairdner, 1915, p. 252).
Since its habits are "largely nocturnal and arboreal, the Bear-cat
is not easily obtained" (Kloss, 1917, p. 294) .
Malay Peninsula. "The Bear-cat ... is generally obtained in
Malacca, and is sometimes kept as a pet. It is easily domesticated,
and becomes very affectionate, and will follow its master like a
dog. It feeds on fruit, also taking small birds." (Ridley, 1895, p. 93.)
"This delightful animal is apparently not uncommon on the main-
land, but I have not heard of it occurring wild in Penang or Singa-
pore. In the Museum at Taiping are specimens from Larut and
Kuala Kangsar, Perak. It is represented in the Museum at Kuala
Lumpor, and is said to be common in Selangor." (Flower, 1900,
pp. 330-331.)
In the Malay Peninsula "Arctictis, Hemigalus, Neofelis, . . .
are not in my opinion vanishing forms. . . . The three carnivores
are all rare but it is extremely difficult to estimate their status in a
country covered with jungle. I see no reason why they should be
classed under Vanishing forms' as there is plenty of country suited
to their requirements and they are not systematically hunted. I
prefer to regard them as uncommon animals, rarely collected. Never-
246
EXTINCT AND VANISHING MAMMALS
theless, they are rare enough to be given total protection." (F. N.
Chasen, in litt., March 31, 1937.)
In Malaya "many with whom I talked . . . were insistent that
such animals as the . . . binturong . . . are to-day practically
non-existent. . . .
"It is true that the loris and binturong fetch a high price in the
Chinese market, but they are numerous." (Comyn-Platt, 1937t>,
P- 48.)
FIG. 26. Binturong (Arctictis binturong subsp.)
Sumatra. In the Korinchi region two specimens are recorded
from Sandaran Agong, 2,450 feet (Robinson and Kloss, 1918, p. 11).
F. N. Chasen (in litt., May 5, 1937) considers the Binturong
much more numerous in Sumatra than in the Malay Peninsula.
"Most specimens seen in captivity originate on the east coast of
Sumatra." It "needs protection as the Chinese use it for medicine."
"In Sumatra, the binturong is found occasionally near Selat
Pandjang. It is not often seen in Rokan and Bengkalis." (Heyn-
sius-Viruly and Van Heurn, 1936, p. 63.)
According to Dr. Hagen, the species is rare in Sumatra. This
agrees with information obtained from natives inland from Palem-
bang. During two years' residence in the Ogan Oeloe Subdivision
only one animal was seen in captivity. (Coomans de Ruiter, 1932,
p. 53.)
Lyon (1908, p. 652) records specimens from Aru Bay, Sungei
Mundau, Siak River, Pulo Payong, and Pulo Tebing Tinggi, eastern
ORDER CARNIVORA: CARNIVORES 247
Sumatra. Pocock (1933, pp. 1018-1019) records additional speci-
mens from Ulu If ok, Perak; Wellesley Province, Straits Settlements;
Tenasserim Village; and Sanderan Agong, Sumatra.
Nias Binturong
ARCTICTIS BINTURONG NIASENSIS Lyon
Arctictis miasensis Lyon, Proc. U. S. Nat. Mus., vol. 52, p. 443, 1916. (Near
Fadoro, Nias Island, off west coast of Sumatra.)
Forty years ago this Binturong was "said not to be common"
(W. L. Abbott, in Lyon, 1916, p. 443) .
"Upper parts of back of head, neck, body, and all of tail, brownish
black, coarsely and rather sparsely grizzled with ochraceous tawny
on the lower back, sides, outerside of legs, and proximal two-thirds
of tail; under parts tawny ochraceous." Tail, 540 mm. The ochra-
ceous-tawny in the present form is replaced by buff or ochraceous-
buff in A. b. binturong. (Lyon, 1916, p. 443.) Cranial and dental
differences are discussed by Miller (1942, pp. 123-124).
This Binturong is apparently restricted to Nias Island. Since the
type specimen was described, a second individual has been recorded
by Miller (1942, pp. 123-124) , who quotes the collector, Frederick A.
Ulmer, Jr., as follows: "The young binturong was purchased alive
from the natives of Soliga in Central Nias and was the only one I
saw, although I heard of one other specimen in captivity near
Gunong Sitoli." Ulmer also refers to it as a "rare animal."
Banka Binturong
ARCTICTIS BINTURONG KERKHOVENI Sody
Arctictis binturong kerkhoveni Sody, Natuurk. Tijdschr. Nederl. Indie, vol. 96,
no. 1, p. 43, 1936. ("Banka Island," Malay Archipelago.)
This form is based upon a tingle specimen from Banka Island.
It is the smallest subspecies. The fur is black, with short buffy
tips to the hairs in some places, especially the head and forelegs;
tail plain black, with light buffy bases to hairs on basal two-thirds
of the ventral surface. Head and body, 600 mm.; tail, 520 mm.
(Sody, 1936, p. 43.)
No information is at hand concerning the frequency of the Bin-
turong on Banka.
Himalayan Binturong
ARCTICTIS BINTURONG ALBIFRONS (F. Cuvier)
Paradoxurus albifrons F. Cuvier, Mem. Mus. Hist. Nat. [Parisl, vol. 9, p. 48,
pi. 4, upper fig., 1822. (Based upon a menagerie specimen in Bengal,
said to have come from Bhutan.)
248 EXTINCT AND VANISHING MAMMALS
FIGS.: Wolf, 1867, pi. 10; Royal Nat. Hist., vol. 1, p. 463, 1893-94; Jour.
Bombay Soc. Nat. Hist., vol. 38, no. 2, suppl., pi. 60, 1935.
This form ranges through Nepal, Sikkim, Bhutan, Assam, Upper
Burma, and Tonkin.
It is distinguished from other forms by the length of its winter
coat and the abundance of long underwool; the color of the body
varies from jet black to tawny'or gray (Pocock, 1933, p. 1030).
"The Binturong appears to be rare in Northern India." Speci-
mens are recorded from Assam; from Endwagyi Lake, in Myitkyina,
Upper Burma; from Fouine, Tonkin; and from Lower Laos. Only
one skull appears to be known. (Pocock, 1933, pp. 1016-1017, 1030.)
"Mr. W. L. Sclater says that it is found even as far west as Simla"
(Lydekker, 1900, p. 334).
Javanese Binturong
ARCTICTIS BINTURONG PENICILLATUS Temminck
(Original reference not found; not in Sherborn; cf. Temminck, Monographies
Mammalogie, vol. 2, p. 310, 1841. Possibly the name was first published by
Temminck sometime prior to 1825 in the prospectus of his "Monographies"
(cf. Valenciennes, 1825, p. 57, footnote). Pocock's citation (1933, p. 1031),
"Temminck, Mon. Mamm. ii. p. 18, 1835," is evidently erroneous; he
gives "Java" as the type locality.)
FIGS.: Ann. Sci. Nat., vol. 4, pi. 1, 1825; Coomans de Ruiter, 1932, p. 54, fig. 12.
The range of this form is Java and Borneo.
It is distinguished from A. b. binturong by its smaller skull and
generally lighter color; pale annulation of the hairs extensive and
profuse (Pocock, 1933, p. 1031). Head and body, 960 mm.; tail,
890 mm. (Schwarz, 1911, p. 636) .
Java. Temminck (1841, vol. 2, p. 311) was inclined to consider
the Binturong the rarest of the mammals of Java and Sumatra. The
species is "apparently rare in Java" (Shortridge, in Thomas and
Wroughton, 1909, p. 386).
Heynsius-Viruly and Van Heurn write (1936, p. 63) :
Few data were received concerning this animal. It was observed in Java
in the vicinity of Madjalengka and near the border of Tomo ....
The binturong is often seen in the Midangan mountains, but it is much
scarcer on the Andjasmoro. None were seen there for the past two years. . . .
Owing to his size, his striking color, his pretty fur and his rather sluggish
motions, the binturong is doomed to be soon exterminated, unless very stern
measures are taken. It occurs on all the Greater Sunda Islands, but is nowhere
common. . . . Young animals become very tame and affectionate in captivity.
This pretty animal . . . should be intensively protected.
Borneo. "These animals are common in parts of Borneo, usually
living in the dense forest, but when in search of fruit they will often
visit gardens" (Hose, 1893, p. 24) .
ORDER CARNIVORA: CARNIVORES 249
Schwarz (1911, p. 636) mentions specimens from Sandakan and
La Datu, North Borneo, and from Sarawak. Pocock (1933, p. 1031)
records others from Mount Mulu, Mount Dulit, and Saribas in
northern Borneo. In the Western Division of Borneo the Binturong
is not rare, and specimens are frequently kept in captivity (Coomans
de Ruiter, 1932, p. 54).
Palawan Binturong
ARCTICTIS BINTURONG WHITEI J. A. Allen
Arctitis [sic] whitei J. A. Allen, Bull. Am. Mus. Nat. Hist., vol. 28, p. 15,
1910. ("Iwahig, Palawan, Philippine Islands")
This form is apparently restricted to Palawan.
It is closely related to A. b. penicillatus, but has a smaller skull;
general color black, strongly washed with fulvous; hairs of nose
and facial region tipped with whitish; neck all round and terminal
fifth of tail black; ears narrowly bordered with white or yellowish.
Head and body, 700 mm.; tail, 610 mm. (J. A. Allen, 1910, p. 15.)
Only about four specimens of this form seem to be on record
(Taylor, 1934, p. 357).
Fossane; Lesser Fossa
FOSSA FOSSA (Schreber)
Viverra Fossa Schreber, Saugthiere, vol. 3, pi. 114, 1776, and p. 424, 1777.
(Based upon "la Fossane" of Buffon (Hist. Nat., vol. 13, p. 163, pi. 20,
1765); type locality, "Madagascar.")
SYNONYM: Fossa daubentonii Gray (1865).
FIGS.: Buffon, op. tit., pi. 20; Schreber, op. cit., pi. 114; J. E. Gray, 1873, pi. 74.
This endemic viverrid of Madagascar is accorded special pro-
tection as a Class A species under the London Convention of 1933.
The ground color is light ashy gray, slightly washed with red-
dish; stripes and spots blackish brown; breast, belly, and legs gray;
tail gray, incompletely ringed with brown (Schreber, 1777, p. 424).
Gray (1873, p. 872) gives the following description: "Brown or red-
dish, closely grizzled with an abundance of white hairs, with four
rows of more or less confluent black spots on each side of the
back, a few black spots on the hinder thighs. The chin, neck, and
belly whitish, more or less obscurely spotted." Daubenton (in
Buffon, 1765, p. 166) gives the following measurements: head and
body, 17 inches; tail, 8J inches.
Our information concerning this animal is very meager. Accord-
ing to Buffon (1765, p. 164), in captivity it eats flesh and fruit and
is especially fond of bananas.
The Mission Zoologique Franco-Anglo-Americaine of 1929-1931
obtained 13 specimens (Delacour, 1932, p. 220).
250 EXTINCT AND VANISHING MAMMALS
"The striped civet of the humid forest is apparently entirely
nocturnal .... Though fairly common" 20 kilometers west of
Vondrozo, "as we found by trapping, none was seen in the daytime.
"Three stomachs from near Vondrozo and one from near Maro-
antsetra all contained insect matter and one contained also a lizard
.... The striped civet was known as 'fanaloka' amongst the Ata-
moor in the southeast." (Rand, 1935, p. 93.)
Family PROTELIDAE: Aard-wolves
The single genus of this family contains one species, which has
been divided into half a dozen subspecies. They range over southern
and eastern Africa. All come within the scope of this work.
Aard-wolf. Maanhaar Jackal (Boer). Faux-loup; Loup de
terre (Fr.). Zibethyane (Ger.)
PROTELES CRISTATUS (Sparrman)
Viverra cristata Sparrman, Resa till Goda Hopps-Udden, vol. 1, p. 581, 1783.
(Near Little Fish River, Somerset East, Cape Province.)
FIGS.: Cuvier, Regne animal, disciples' ed., Mamm., atlas, pi. 40, fig. 3,
1836-1849; Proc. Zool. Soc. London 1869, pi. 36; W. L. Sclater, 1900,
vol. 1, p. 81, fig. 21; Anderson and de Winton, 1902, pi. 28; Derscheid,
1925, pi. A; Pocock, 1937, p. 758, fig.
The Aard-wolf is of particular scientific interest as the sole repre-
sentative of the family Protelidae; and it has been accorded rigid
protection as a Class A mammal under the London Convention of
1933.
Six subspecies have been proposed, as listed below, but since their
distributional limits have not been worked out, all will be included
in this account of the species as a whole.
Proteles cristatus cristatus (Sparrman) . Cape Aard-wolf. (Type
locality as given above.)
Form hyenalike; general color dirty yellowish gray, with project-
ing coarse hairs, black and white; an erectile black mane from
nape to tail; seven to nine transverse black stripes on sides; upper
parts of limbs with indistinct black bands; feet black; face, lower
jaws, and chin brown; tail yellowish at base, rest black. Head and
body, 32 inches; tail without hairs, 6 inches. (W. L. Sclater, 1900,
vol. 1, pp. 80-81.)
Proteles cristatus pallidior Cabrera, Ann. Mag. Nat. Hist., ser. 8,
vol. 6, p. 464, 1910. Nubian Aard-wolf. ("Suakim," Anglo-Egyptian
Sudan.)
General color pale yellowish cream ; body hairs unicolored ; cheeks
and sides of neck not rufous as in true cristatus; mane with very
little black; feet brownish, not black; tail black only at tip (Cabrera,
1910, p. 464).
ORDER CARNIVORA: CARNIVORES 251
Proteles cristatus septentrionalis Rothschild, Novit. Zool., vol. 9,
p. 443, 1902. Somali Aard-wolf. ("Somaliland.")
Creamy white, washed with buff on neck and sides of rump;
stripes less defined than in cristatus; mane black, variegated with
creamy white (W. Rothschild, 1902, p. 443).
Proteles cristatus termes Heller, Smithsonian Misc. Coll., vol. 61,
no. 13, p. 9, 1913. Masailand Aard-wolf. ("Headwaters of the
Amala River west of the Loita Plains, British East Africa.")
Interorbital region black, crown grizzled, feet black, and ears
blackish as in cristatus; body stripes narrow; ground color grayish
fulvous; tail black on terminal third; muzzle and chin black; throat
grayish buffy. Head and body, 680 mm.; tail, 310 mm. (E. Heller,
19136, p. 9).
Proteles cristatus transvaalensis Roberts, Ann. Transvaal Mus.,
vol. 15, pt. 1, p. 6, 1932. Transvaal Aard-wolf. ("Roodekuil, Pre-
toria," Transvaal.)
Pale buffy; face, bands on limbs and body, and distal third of
tail dark brown; mane rufous-white, the hairs with three brown
bands and blackish tips. Head and body, 650 mm.; tail,, 270 mm.
(Roberts, 1932, p. 6.)
Proteles cristatus harrisoni Rothschild, Novit. Zool., vol. 9, p. 443,
1902. Angola Aard-wolf. ("Umpata, Mossamedes district, S. An-
gola.")
Head white, grizzled with black; body pale orange rufous; stripes
less developed than in other races; mane and tail black, variegated
with rufous (Rothschild, 1902, p. 443) .
The species as a whole has a wide range over southern and eastern
Africa, from Cape Province and Natal north to Angola, Northern
Rhodesia, and Anglo-Egyptian Sudan. It is apparently absent
along the low-lying east coast from the Transkei district of Cape
Province to Portuguese East Africa. (Shortridge, 1934, vol. 1,
p. 150.)
Anglo-Egyptian Sudan. Specimens have been recorded from the
vicinity of Suakin (Anderson and de Winton, 1902, p. 198; Cabrera,
1910, p. 465) and from the Blue Nile (Butler, in Maydon, 1932,
p. 151).
Eritrea. This animal is rather widespread, if not common, in the
greater part of the country, especially in the southwest (Zammarano,
1930, p. 77).
Ethiopia. It is "common on the lowlands and foot hills wherever
white ants abound" (Wylde, 1901, p. 485).
British Somaliland. "The aardwolf is found sparsely scattered
throughout Somaliland. They are almost invariably seen alone."
(Drake-Brockman, 1910, p. 38.)
252 EXTINCT AND VANISHING MAMMALS
Italian Somaliland. Recorded by De Beaux (1935, p. 12).
Kenya. Hollister (1918, pt. 1, p. 138) lists specimens from the
following localities: Kabalolot Hill, Sotik; Northern Guaso Nyiro
River; Telek River, Sotik; and Ulukenia Hills.
Tanganyika Territory. Recorded from Tabora (Matschie, 1895,
p. 62).
Northern Rhodesia. "I am reliably informed that specimens
have been obtained at Tara and Kalomo in the Batoka Province.
Elsewhere the natives do not seem to have heard of it." (Pitman,
1934, p. 162.)
Southern Rhodesia. It ranges rather sparsely over this country
(western Matabeleland, etc.) (Shortridge, 1934, vol. 1, p. 150).
Transvaal. "In the Eastern Transvaal the Aard Wolf is not
found in the low-veld proper; it occurs in the more open country
among the foothills of the Drakensberg at a height of over 1,500
feet" (Hamilton, in Shortrklge, 1934, vol. 1, p. 150). A specimen is
recorded from Potchefstrom (W. L. Sclater, 1900, vol. 1, p. 82).
Natal. "It is fairly common in Natal" (Warren, in Shortridge,
1934, vol. 1, p. 150).
Cape Province. The Aard-wolf is reported as not uncommon
throughout the colony (W. L. Sclater, 1900, vol. 1, p. 81).
Bechuanaland. "The Kalahari Sand-Plains" are "perhaps the
regions in which it is most plentiful. . . . The karross-making
tribes in Bechuanaland . . . are said to procure most of their aard
wolf skins with the aid of dogs." (Shortridge, 1934, vol. 1, p. 150.)
As many as 14 have been seen together in the Kalahari (Langdon,
in Shortridge, 1934, vol. 1, p. 151).
South-West Africa. "Proteles is widely distributed throughout
South -West Africa; nowhere very abundantly. It is apparently
rather scarce along the valley of the Orange River, and northwards
in the neighbourhood of the Okavango and in the Caprivi. . . .
"The Aard Wolf is fairly plentiful around Gobabis and in the
sand-plains generally; and is also familiar in Namaqualand, Da-
maraland, the Kaokoveld, Ovamboland, and the Namutoni Game
Reserve." (Shortridge, 1934, vol. 1, pp. 149-150.)
Angola. It is rather common in the south of Angola but much
rarer in the north. A skin was brought in to Vila da Ponte, where
the animal was unknown to the natives. (Monard, 1931, p. 66, and
1935, p. 228.) The type locality of the subspecies harrisoni is Um-
pata in the Mossamedes district.
Economic status. "The coat is very handsome, and ... its skin
is more sought after [than that of the hyenas] by some of the native
tribes notably the Bechuanas, who hunt and trap it systematically"
(Bryden, 1899, p. 599).
"Sparrman and other authors who have examined the stomachs
ORDER CARNIVORA: CARNIVORES
253
of these animals, found that they contain nothing but termites or
white ants; this is further confirmed by Mr. Cloete, who writes that
he has examined the stomachs of more than fifty, and never found
any trace of anything else than a purely insectivorous diet, ants
being the chief constituent." However, farmers report that this
species kills kids and lambs merely for the sake of the milk con-
tained in their stomachs. (W. L. Sclater, 1900, vol. 1, p. 82.)
The food consists of insects particularly termites, locusts, beetles,
and grubs. The animal has been accused of killing lambs and kids,
Fie. 27. Aard-wolf (Proteles cristatus subsp.)
but evidently without justification and through confusion with the
Jackal. Its weak dentition is sufficient evidence of its harmlessness
in respect to livestock. Yet, despite its inoffensive nature and the
distinct service it performs in destroying such agricultural pests as
termites, it was, in former years, officially listed by several govern-
ments as "harmful," and a bounty of half a pound sterling was paid
in the Cape Province for each Aard-wolf killed. (Derscheid, 1925,
P. [78].)
It feeds to some extent on "small rodents, reptiles, and the nest-
lings and eggs of ground-nesting birds" (Shortridge, 1934, vol. 1,
p. 151).
In South Africa generally it is subjected to a great deal of irre-
sponsible persecution, and is becoming scarce in farming and other
settled districts. However, it is in no immediate danger of actual
extermination. (G. C. Shortridge, in litt., October 14, 1937.)
254 EXTINCT AND VANISHING MAMMALS
Although normally the animal has no unpleasant odor, it is able
to eject an evil-smelling fluid from its anal glands as a defense
against such enemies as dogs.
Family FELIDAE: Cats
This family is nearly cosmopolitan, but it does not occur natur-
ally in Greenland, the eastern Malay Archipelago, or Australasia.
Twenty or more genera are recognized by some authorities, and
there are probably more than 250 forms. Dr. Allen discusses 12
North American forms in the preceding volume (1942), while 27
Old World forms are dealt with in the following pages. Man's
prejudice against some of the larger members of the cat family
(such as Lions, Tigers, Leopards, and Cougars) is linked with his
necessary efforts to defend himself or his livestock from their attacks.
Fossa
CRYPTOPROCTA FEROX Bennett
Cryptoprocta jerox Bennett, Proc. Zool. Soc. London 1833, p. 46, 1833. ("Mada-
gascar.")
FIGS.: Bennett, 1834, pi. 21; Schreber, Saugthiere, suppl. vol. 2, pi. 125CC,
1841; Schlegel and Pollen, 1868, pi. 8; Proc. Zool. Soc. London 1895, pi.
26; Beddard, 1902, p. 405, fig. 199; Elliot, 1907, p. 397, fig. 43; Kaudern,
1915, pi. 3, fig. 2; Sibree, 1915, pi. facing p. 302; Pocock, 1937, p. 760, fig.
This largest carnivore of Madagascar occurs in limited numbers
and will probably require protection if it is to survive in the
dwindling forests of that great island. It is an endemic species.
Since Bennett's type specimen was distinctly immature, the fol-
lowing description of an adult male is derived from Schlegel and
Pollen (1868, pp. 13-14). Hairs of upper parts ringed with brown
and pale reddish yellow; lower parts of head and body uniform
reddish yellow, taking on a strong rusty tint toward the middle of
the venter. Total length, 56 inches; tail, 26 inches.
Bennett (1834, p. 140) quotes Charles Telfair as follows: "It is
the most savage creature of its size I ever met with: its motions
and power and activity were those of a tiger: and it had the
same appetites for blood and destruction of animal life."
Milne Edwards and Grandidier write (1867, p. 317) that it occurs
rather commonly on the west coast, from the River Mangouke
[Mangoky] northward. Three specimens were secured between
Morondava and Manharrive [Maharivo?]. The animal often carries
off goats and especially kids.
This animal is very carnivorous and is endowed with great
strength. It is dangerous to man only when wounded or in rut.
At other times "it steadily flees from man. At the mating season
ORDER CARNIVORA: CARNIVORES 255
it is often seen in bands of four to eight individuals. It is said to be
fond of lemurs and to pursue them in trees. It is also destructive
to poultry, young pigs, and other domestic animals. The natives
really fear this species, but they enjoy its flesh. (Schlegel and
Pollen, 1868, pp. 15-16.)
According to Milne Edwards and Grandidier (1875a, p. 341, foot-
note) , this is the only native animal of Madagascar that the Saka-
lava (a western tribe) have been able sometimes, but very rarely,
to train for hunting the Wild Hog (Potamochoerus larvatus) .
Kaudern states (1915, pp. 79-80) that Cryptoprocta appears to
be distributed over the entire island and that it is probably nowhere
FIG. 28. Fossa (Cryptoprocta jerox). After photo in Brehm.
rare. In northwestern Madagascar it was very common. He saw
the animal three times in the wild at Ste. Marie de Marovoay on
the Betsiboka River, and its tracks were observed everywhere in
the sand. One was killed there in a poultry yard. Another was
secured at Katsepe on the Bay of Bombetoke, and two live young
ones were brought in by natives at Andranolava, in north central
Madagascar. Black individuals are reported from the interior and
from the great rain forests on the east coast.
According to Sibree (1915, pp. 302-303), the northwest coast is
the animal's "special habitat. This creature is called by the people,
Fosa . . . , and although small is very ferocious .... Examples
of the fosa have been seen in the outskirts of the upper belt of forest
on the east side of the island .... A specimen I once saw was of a
beautiful black colour, but I believe this was only a variety, and
not a distinct species from the brown animal. The fosa is much
dreaded by the Malagasy, and, from its mode of attack, appears to
be like an immense weasel, attacking large animals, such as the
wild boar and even oxen."
G. M. Allen (1918, p. 514) records a specimen from the vicinity
of Tulear.
256 EXTINCT AND VANISHING MAMMALS
Petit (1931, p. 588) records a female and its three young ones
captured in 1922 in the region of Tamatave, and two young ones
taken in the region of Fenerive on the east coast.
The Mission Zoologique Franco-Anglo-Americaine of 1929-1931
secured 6 specimens of this species, as compared with 13 specimens
of Fossa jossa (Delacour, 1932, p. 220).
Rand (1935, pp. 93-94) says:
The fossa inhabits the rain forest of the east and the dryer forest of the
west at least as far south as Tabiky [inland from Cape St. Vincent], and was
well known to the natives. [Two were seen near Tsarakibany and Maromandia
during the daytime, though the natives said it was nocturnal.] This viverrid
was much disliked by the natives because of its raids on their fowls. Twice
I saw fossa skins in the possession of natives, but this was probably due to
European influence as the natives rarely use mammal skins for any purpose.
One large fossa was brought to me that had been run down with dogs and
speared. From the natives we heard no accounts of its attacking sheep or
young cattle and its reputation in literature for ferocity and the fear with
which it is regarded by the natives is exaggeration. My gun boy had a
particular antipathy for it because, he said, in his country near Vondrozo,
where the dead are walled up in caves, the fossa sometimes dug out the
corpses and fed on them. The natives universally called it "fossa."
European Wildcat. Chat sauvage (Fr.). Wildkatze (Ger.).
Gato monies; Gato salvage (Sp.) Gatto selvatico (It.)
FELIS SILVESTRIS SILVESTRIS Schreber
Felis (Catus) silvestris Schreber, Saugthiere, vol. 3, p. 397, pis. 107A, 107 Aa,
1777. (Germany.)
FIGS.: Gervais, Hist. Nat. Mammif., pt. 2, pi. 17, 1855; Blasius, 1857, p. 162,
fig. 101; Elliot, 1883, pi. 30; Royal Nat. Hist., vol. 1, pi. facing p. 422,
1893-94; Hamilton, 1896, frontisp.; Martin, 1910, pi. 27; Cabrera, 1914,
pi. 9, fig. 1; Zeitschr. f. Saugetierk., vol. 7, pi. 7, fig. 7, 1932; Colosi, 1933,
p. 55, fig.; Didier and Rode, 1935, p. 283, fig. 163; Schmidt, 1938, pi. 5.
The typical European Wildcat shows a very general and marked
recession in France and central Europe, amounting to extirpation
in many parts of its former range. Apparently its chief remaining
stronghold is in the Balkan countries.
More definite light is needed on the question as to whether inter-
breeding with feral Domestic Cats takes place at all or on a
sufficient scale to menace the Wildcat's survival as a pure-bred
species. Fatio states (1869, p. 276) that hybrids are sometimes
met with in Switzerland, and that he has examined a number of
specimens; the pelage, he adds, is often spotted with white. Ferrant
(1931, p. 62), in discussing the Wildcat in Luxembourg, says that it
mates frequently with feral Domestic Cats. Prof. M. Hirtz refers
(in litt., December, 1936) to hybrids in Yugoslavia, and the National
Council for Nature Protection does likewise (in litt., October, 1936)
in Poland. On the other hand, Pocock (1907, pp. 165-166) is rather
skeptical in regard to the alleged interbreeding.
ORDER CARNIVORA: CARNIVORES 257
Hamilton speaks rather emphatically in his monograph (1896,
p. iv) : "On a careful examination of a number of examples of the
Wild Cat of the present time I found many indications of a mixture
of the two races.
"It would seem as if the original Wild Cat, as it existed in the
olden days, has been almost exterminated throughout Europe, and
that its place has been taken by a mongrel race, the result of con-
tinual interbreeding during many centuries (2000 years) of the
Wild and the imported Domestic Cat."
The Wildcat is slightly larger than the Domestic Cat; fur longer,
this being especially noticeable in the tail ; general color approaching
the smoke gray of Ridgway; dark markings on sides and legs tend-
ing to be faint, brownish, and ill-defined; tail, abruptly rounded
at the black tip, with two to four more or less complete black rings ;
tip of ear slightly blackish. Head and body, 481-545 mm.; tail,
309-310 mm. (Miller, 1912, pp. 457-463.) Head and body, 450-
700 mm.; tail, 200-300 mm. (Didier and Rode, 1935, p. 284).
The typical subspecies ranges from northern Spain, France, and
northern Germany eastward to Poland and Russia, and southward
to Italy and the Balkan Peninsula.
In Spain it inhabits the Pyrenean, Cantabrian, and north central
districts, and it may extend to northern Portugal. Probably the
Douro and the Ebro constitute its southern limits. (Cabrera, 1914,
p. 204.) "Curiously enough there is no specimen in the Museums
of Ponferrada, Lugo, and Santiago. I was left in doubt as to whether
the wild cat occurs in the Sierra de Picos, in Galicia, or in the Picos
de Europa." (Gadow, 1897, p. 367.)
In France, where it was formerly very common, the Wildcat is
at present in the process of disappearing, but may still be met with
in certain regions, such as the Ardennes, the Forest of Orleans, the
Pyrenees-Orientales, and the Forest of Carnelle. It preys upon
rabbits, hares, squirrels, rats, young Roebucks and Wild Boars,
grouse, partridges, and pheasants, and even eats fishes. It is a very
injurious animal, which one need not hesitate to destroy. (Didier
and Rode, 1935, p. 287.)
Hunting has contributed in part to its depletion in France, but
does not explain this altogether. The species is very much in danger,
and we do not think that protective measures can be effective. (E.
Bourdelle, in litt., March 6, 1937.)
In Belgium the species has become very rare, but still exists in
some forests in the Province of Luxembourg. It is systematically
destroyed as an injurious animal, while another cause of depletion
is deforestation. Total protection ought to be adopted. (Musee
Royale d'Histoire Naturelle de Belgique, in litt., September, 1936.)
258 EXTINCT AND VANISHING MAMMALS
In the Grand Duchy of Luxembourg the Wildcat is rather com-
mon in the extensive forests of the Ardennes, in the environs of
Echternach, Grevenmacher, Manternach, and Fischbach, in the
Grunewald, etc. (Ferrant, 1931, p. 61).
In The Netherlands the species was long since exterminated (Van
den Brink, 1931, p. 174).
In Germany the Wildcat has survived better than the larger
carnivores; it occurs in very small numbers in the Bavarian moun-
tains, the Black Forest, the Odenwald, and the Riesengebirge. From
1850 to 1860 ten animals were killed in Gotha; in 1885-86, two in
Silesia; in 1928 an unquestionably pure-blooded male was taken in
the Harz Mountains, and in the same year a male in the Kurische
Nehrung. (Krumbiegel, 1930, pp. 5-6.) The Wildcat is still regu-
larly observed in the Eifel, in the Moselle Mountains, and in the
Hunsriick, and there is one from the Pfalz in the Koln Zoological
Garden (Hauchecorne, Zeitschr. f. Saugetierk., vol. 9, p. 4, 1934).
The animal is almost exterminated in Germany, and is protected
as a natural monument (Internationale Gesellschaft zur Erhaltung
des Wisents, in litt., October, 1936) .
In Denmark bones of the Wildcat have been found in kitchen
middens, but there is no record within historical times (Winge,
1908, p. 116).
In Switzerland it appears to have been abundant in the sixteenth
century and was then the object of much hunting; but it had
become rare by the beginning of the nineteenth century. In the
1860's some were killed each year in the Alps and the Jura, and a
few were still found in the cantons of Bern, Lucerne, Unterwalden,
Uri, Schwyz, Glarus, Thiirgau, and Valais. The most were found in
the Jura region, from Geneva to Basle. The species seemed to have
disappeared from Ticino. (Fatio, 1869, p. 275.) During the last
decades it has become very rare and is probably extinct, although
it is possible that a few survive in the forests of the Alps and the
western Jura (Federal Forest, Game, and Fish Inspection, Bern,
in- litt., March, 1937) .
Wildcats have almost disappeared from northern Italy, and are
rare everywhere except in the Maremma, in the southern provinces
of Gargano, and in Calabria (Colosi, 1933, p. 56). [The animal of
the Tuscan Maremma is regarded by Martorelli (1896, p. 266) as
identical with the Sardinian Wildcat (Felis sarda).] The Wildcat
still occurs in the Sila Mountains of Calabria (Hecht, 1932, p. 23).
According to the Laboratorio di Zoologia Applicata a Caccia (in
litt., September, 1936) , the animal is scattered through Sicily as well
as the Italian Peninsula ; in legislation it is rated as a harmful species.
In former times it was probably found everywhere in Austria.
It is now exterminated in Burgenland but is said to survive in
ORDER CARNIVORA: CARNIVORES 259
Rosenbachtal in Carinthia. In 1926 two specimens were killed in
the district of Volkermarkt, Carinthia. In Lower Austria the last
one was killed in 1912. The species is no longer found in Salzburg.
In the Tyrol 26 specimens are said to have been shot in 1876. By
1888 it was almost exterminated in northern Tyrol, but was con-
sidered more frequent in southern Tyrol. In Vorarlberg it was
reported as late as 1918, but has now disappeared. (G. Schlesinger,
in Hit., March, 1937.)
Within the boundaries of the present Hungary it was common
before World War I, but is now decreasing. It has no legal pro-
tection. It is also found in the northern part of the former Hungary
(now Czechoslovakia) , but not together with the Lynx. ( J. Schenk,
in Hit., November, 1936.)
In comparison with its status in most other parts of Europe,
the Wildcat is comparatively common in Yugoslavia. Considerable
numbers are found only in certain regions, and especially in the
enormous oak forests of Slavonia. The statistics are uncertain,
since they probably include hybrids and feral Domestic Cats. The
reported annual kill from 1891 to 1921 was about 500 to 1,000
specimens, reaching a maximum of 1,207 in 1904. The minimum
kills were 420 in 1918 and 331 in 1931. These figures pertain chiefly
to Croatia and Slavonia. (M. Hirtz, in litt., December, 1936.) In
northwestern Croatia the animal is rare near Jasenak and near
Otocac (Wettstein, 1928, p. 35).
Lord Lilford wrote of frequently meeting with Wildcats in the
Province of Epirus, near the boundary between Albania and Greece
(Hamilton, 1896, p. 35).
The Wildcat is found throughout Greece except on the islands.
A decrease has been observed, but there is no danger of extermina-
tion. (Game Department, Ministry of Agriculture, Greece, in litt.,
October, 1936.) (The Wildcat of the Peloponnesus is discussed on
a subsequent page, under the name of Felis silvestris morea.)
Turkey is included in the range by Blasius (1857, p. 166).
In Bulgaria the Wildcat occurs in considerable numbers and is
in no danger of extermination. The skins are marketed locally.
(Bulgarian Game Association "Sokol," in litt., February, 1937.)
H. W. Shoemaker (in litt., June 30, 1932) is of the opinion that
the Wildcats in this country are being rapidly destroyed.
The species is common in Rumania, occurring in nearly all forests
of greater or lesser altitude; it is also frequently found in the
lowland forests (District of Ilfoo) and in the flood lands of the
Danube (Calinescu, 1930, p. 366).
In Poland it occurred formerly from the Carpathians to the
Baltic but is now met with chiefly in the eastern Carpathians; it
is also found in the forests along the Dniester (Zurawno, Stanis-
260 EXTINCT AND VANISHING MAMMALS
lawow) and in Podolia. It is nowhere common (Niezabitowski,
1934, pp. 190-191). By 1936 the species was considered restricted
to the Carpathians, where its numbers are roughly estimated at 300.
It is protected from February 1 to September 30. (M. Siedlecki
and National Council for Nature Protection, Poland, in litt., October,
1936.) The proposed International Tatra Park and the proposed
International Park of Poland, Czechoslovakia, and Rumania will, it
is hoped, provide absolute protection for the Wildcat and other
species (Benedyct Fulinski, MS., 1933).
The following summary of the Wildcat's status in Russia is de-
rived from Ognev (Ogneff, 1930, pp. 55-58) . The information is frag-
mentary and suffers from lack of material. Pallas (1811-1831) de-
nied the animal's existence throughout Russia except in the Caucasus.
Georgi (1800) reported it in the southwestern governments, on the
Dniester, and in the central Urals (Bashkiri). Brandt (1853) be-
lieved in an early, much wider distribution, as far as the central gov-
ernments and perhaps to the Urals. Kessler (1856, 1858) records the
Wildcat in Volhynia and Podolia. In 1854 it was reported in the
Governments of Grodno, Vitebsk, and Kovno. Sabaneeff (1878)
considered the Government of Minsk the center of its distribution
in western Russia. According to Charlemagne (1920), it is now
very rare in Volhynia and in the vicinity of Odessa and Tiraspol
(Government of Kherson). Old reports from central and northern
Russia in the latter part of the past century may have been based
upon feral Domestic Cats. The question as to whether the Wildcat
ever occurred in the Urals is unsolved.
On the subject of general depletion Elliot comments (1883, text
to pi. 30) : "Various are the causes that have effected this ; probably
the chief one is the constant persecution to which the animal has
been subjected, as this species has but few friends, and no quarter
is shown when it is met with in the forest."
Hamilton (1896, pp. 31, 95) remarks on the spread of the Domestic
Cat with the increase of the human population, and the resulting
interbreeding with the Wildcat, as possibly a chief factor in the
disappearance of the pure-bred wild animal in Europe generally.
British Wildcat
FELIS SILVESTRIS GRAMPIA Miller
Felis grampia Miller, Ann. Mag. Nat. Hist., ser. 7, vol. 20, p. 396, 1907.
("Invermoriston District, Inverness, Scotland.")
Fios.: Millais, 1904, pis. facing pp. 166, 170, 172, 174, 178; Pocock, 1937,
p. 777, fig.
The British Wildcat formerly occurred throughout Great Britain
but is now restricted to the wilder portions of Scotland (Miller,
1912, p. 464).
ORDER CARNIVORA: CARNIVORES 261
It is like Felis silvestris silvestris of continental Europe, but the
general color is darker, approaching broccoli-brown; dark markings
on sides and legs tending to be extensive, blackish, and well defined ;
upper side of feet and inner surface of hind legs ochraceous-buff,
under side of body duller; intercrural and pectoral white areas well
defined; middle of chest mottled with black; dark markings on tail,
legs, and upper parts similar to, but more definite than, those of
F. s. silvestris (Miller, 1912, p. 464). Males: head and body, 558-
660 mm. ; tail, 280-355 mm. Females slightly smaller. (Millais, 1904,
p. 170.)
The following account is condensed from Millais (1904, pp. 170-
180) . It is not known when the Wildcat became extinct in southern
and central England, but it probably lingered until the forests were
cleared. In Wales it may have survived till about the end of the
nineteenth century. Approximate dates of last records in England
are: Yorkshire, 1840; Lake District, 1843. In churchwardens' ac-
counts and other records there is mention of bounties paid for Wild-
cats in the sixteenth, seventeenth, eighteenth, and even nineteenth
centuries.
Last dates in certain counties of Scotland are: Berwickshire,
1849; Dumfriesshire, Wigtown, and Kirkcudbright, about 1832;
Dumbarton, 1857; Perthshire, 1870-71; Aberdeenshire, 1891; Forfar,
Kincardine, Banff, Elgin, and Nairn, practically extinct since 1850.
"Northern and western Inverness is, with western Ross-shire, the
main stronghold of the Wild Cat to-day." Up to 1904 William
Macleay, of Inverness, annually received eight or ten specimens,
chiefly from Glenmoriston and Balmacaan, west of Loch Ness. "In
Sutherland the Duke of Sutherland does not allow the slaughter of
Wild Cats to take place," and the animals are on the increase in
certain parts. In Caithness the species was never common but was
reported as occasional until 1845, and it evidently survived to a
somewhat later date.
The Wildcat's prey includes poultry, lambs, and roe fawns. But
since it keeps down certain animals such as grouse, hares, and
rabbits that are considered undesirable in deer forests, it receives
protection from the sportsmen who control these forests.
According to the minutes of a meeting of the Society for the
Preservation of the Fauna of the Empire in October, 1922, "a cir-
cular to owners and tenants of deer forests in Scotland, asking them
to protect wild cats and martens, had been well received." Several
years later it is reported that "we get very gratifying reports in
regard to the preservation by land owners of wild cats and pole cats"
(Onslow, 1929, p. 7).
"Mr. N. B. Kinnear remarked that the wild cat was now not rare
in the north of Scotland, and a good account of its increase and
262 EXTINCT AND VANISHING MAMMALS
spread was given by the late Mr. J. G. Millais in 'The Times' of 26
October 1926. According to that article the wild cat reappeared in
Inverness-shire, from north of the Caledonian Canal, about 1912
and, after becoming established round Lochs Ericht and Laggan,
spread farther south into Perthshire, where one was killed at
Murthly, twelve miles from Perth, in 1925.
"Mr. Kinnear further stated that owing to the increase of tree
planting the safety of the wild cat appeared to be assured, as it had
taken to the young plantations on account of the rabbits and, where
the plantations were under the charge of the Forestry Commission,
the cats were encouraged, as they helped to keep down the rabbits."
(Kinnear, 1934, p. 68.)
"The War granted a respite to the Carnivora, and the Wild Cat,
which but for that event would probably have been exterminated by
now, increased in numbers. But now the persecution of this animal
and other carnivores is in full swing again; and apart from a slight
possibility of help coming from the third cause of change, dealt
with below [planting of thousands of acres in the Highlands with
conifers], it is probable that the Wild Cat will be brought to the
verge of extinction again before long." (Hinton, 1935, pp. 33-34.)
Peloponnesian Wildcat
FELLS SILVESTRIS MOREA Trouessart
[Felis catus] morea Trouessart, Cat. Mamm., quinq. suppl., fasc. 1, p. 273,
1904. (Based upon the "Felis catus ferus L. var. e Morea" of Reichenbach,
Vollstandigste Naturgeschichte, Raubsaugethiere, p. 362, 1852, ex Bory
de Saint-Vincent, Exped. Sci. Moree, atlas, ser. 3, zool., pi. 1, A, 1833;
type locality, as restricted by Harper (1940, p. 194), "above Dragomanou,
near Mt. Diaphorti, west central Morea (Peloponnesus), Greece.")
FIGS.: Bory de Saint-Vincent, Exped. Sci. Moree, atlas, ser. 3, zool., pi. 1, A,
1833; Reichenbach, Praktisch-gemeinniitzige Naturgeschichte, Kupfer-
sammlung, pt. 1, Raubsaugthiere, pi. 80, fig. 639, 1837(?).
This form of southern Greece, while evidently less common than
formerly, does not seem to be threatened with extinction and is
included here chiefly for the purpose of rounding out the picture
of the European Wildcats.
It differs from F. s. silvestris in its generally isabelline coloration,
in the absence of distinct stripes on the sides, and in having the
black rings on the tail straight and clearly defined; lateral stripes
replaced by irregular brownish-rufous marblings; feet unspotted
(Trouessart, 1910, p. 100).
Bory de Saint-Vincent states (1836, vol. 1, p. 396) that among
the oak-dotted pastures of the type locality near Mount Diaphorti
the Wildcats occur in larger numbers than elsewhere. Here he
collected the type specimen from the high branches of an oak.
ORDER CARNIVORA: CARNIVORES 263
According to Geoffrey (in Bory de Saint-Vincent, 1833, vol. 3,
pt. 1, zool., p. 13), the animal is very common in certain mountain-
ous parts of Arcadia, especially in the Canton of Karytaena and on
the slopes of Mount Diaphorti. It is destructive to poultry, small
birds and mammals, and partridges.
The Game Department, Greek Ministry of Agriculture, reports
(in litt., October, 1936) a general decrease in the numbers of Wild-
cats in Greece.
[The Spanish Wildcat (Felis silvestris tartessia *) inhabits the
Iberian Peninsula south of the Douro and the Ebro, and still
abounds in the wilder parts. While interbreeding with feral Domestic
Cats probably takes place, no evidence of it has been found. (Ca-
brera, 1914, pp. 205-206.)
The Caucasian Wildcat (Felis silvestris Caucasians 2 ) occurs in
all the mountain forests and in the greater part of the forested
lowlands of the Caucasus region. It is generally reported as very
common. (Ognev, 1930, p. 58.) ]
Cretan Wildcat. Chat sauvage de Crete (Fr.)
FELIS AGRIUS Bate
Felis ocreata agrius Bate, Proc. Zool. Soc. London 1905, pt. 2, p. 317, 1906.
(Type skin bought in the bazaar at Khania, Crete.)
This species, which is confined to the island of Crete, may be in
danger of extinction by "dilution," consisting in this case of inter-
breeding with feral Domestic Cats.
The general color is yellowish gray; no black markings on body
or legs, but indications of brownish shoulder stripes and dorsal
stripe ; tail with black tip and two or three black subterminal rings ;
ear blackish at tip (Miller, 1912, p. 470) .
Raulin (1869, p. 1033) records the species from the woods of the
lower zones.
"Hybrids between F. o. agrius and the domestic cat of the island
appear to be not uncommon, and this can easily be accounted for
by the fact that formerly small villages were often totally deserted
for a considerable time, or possibly entirely, during the insurrections
which occur so frequently in Crete, when the cats, as well as the
villagers, are forced to take to a life in the hills. Skins of these
hybrids, which are generally of large size like the true wild race,
may often be seen hanging up in the bazaars at Khania and Candia."
(Bate, 1906, p. 318.)
^ Felis tartessia Miller, Ann. Mag. Nat. Hist., ser. 7, vol. 20, p. 397, 1907.
("Coto Donana, near Jerez de la Frontera, [Huelva,] Spain.")
2 Felis catus caucasicus Satunin, Mitteil. Kaukas. Mus., vol. 2, pts. 2-4, pp.
154 (Russian) and 316 (German), 1906. (Caucasus region.)
264 EXTINCT AND VANISHING MAMMALS
The same author (in Trevor-Batty e, 1913, p. 255) refers to the
Cretan Wildcat as "not uncommon in the island."
The Game Division, Forestry Department, Greek Ministry of
Agriculture, seems to consider (in Hit., March, 1937) that this animal
is derived from Domestic Cats which the inhabitants left when they
had to flee during the war of independence, and also from those
left by the departing Turks when the exchange of populations took
place after 1922.
It may be added that Pocock (1907, pp. 151, 160) evidently
believes that the type specimen was a Domestic Cat or a feral repre-
sentative of one.
Cor si can Wildcat. Chat sauvage de Corse (Fr.)
FELIS BEYI Lavauden
Felis reyi Lavauden, C. R. Acad. Sci. [Paris], vol. 189, p. 1023, 1929. ("Foret
d'Aunes des bords de la lagune de Biguglia (Sud de Bastia)," Corsica.)
This Wildcat is included because of its interest as an insular form
and because of the generally uncertain future of the Wildcats of
Europe, rather than on account of any definitely recorded decrease.
It is smaller than Felis silvestris silvestris; pelage very dark, with
a darker, rather indistinct dorsal stripe; hind feet with black marks
like those of African Wildcats; back of the ear dark brown. Head
and body, 580 mm. ; tail, 270 mm. Weight, 2 kg. (small females) to
5 kg. (large males). (Lavauden, 1929, pp. 1023-1024.)
No Wildcat had been recorded from Corsica before 1929, and
only three specimens have been studied so far, but the species is
not extremely rare. It is found throughout Corsica in the high
mountains, the forests, the thickets of the hills, and the shrubbery
of the plains. The Corsican hunters do not bother to seek the animal
because of the low value of its fur. (Lavauden, 1929, p. 1024.)
Sardinian Wildcat. Chat sauvage de Sardaigne (Fr.). Gatto
selvatico di Sardegna (It.)
FELIS SARDA Lataste
[Felis libyca] var. sarda Lataste, Act. Soc. Linn. Bordeaux, vol. 39, p. 231,
1885. ("Sarrabus (Sardaigne).")
FIGS.: Martorelli, 1896, pis. 1, 2.
While little information is at hand concerning the numerical
status of this species, it is possibly being subjected, like other
Wildcats of Europe, to the process of extinction by "dilution" in
addition to direct persecution.
It differs from Felis silvestris in its shorter fur and more slender
tail (hairs at middle averaging about 30 mm. instead of 40 mm.) ;
ORDER CARNIVORA: CARNIVORES 265
hairs of median dorsal line slightly elongated and stiffened; dark
markings obsolete, the back and sides grayish or brownish, without
definite stripes; back of ear yellowish clay-color, the tip black; tail
with well-defined black tip. Head and body, 600 mm. ; tail, 300 mm.
(Miller, 1912, pp. 468, 470.)
While this species was originally described from Sardinia, the
same animal is recorded by Martorelli (1896, p. 266) from the
Maremma of Tuscany in western Italy.
It is found throughout Sardinia but is not numerous there, and
in legislation is rated as harmful (Laboratorio di Zoologia Applicata
a Caccia, in litt., September, 1936).
The animal of Tuscany is said to be not rare (Colosi, 1933, p. 56) .
It is considered a fierce destroyer of hares, pheasants, and other
game and consequently is much persecuted (Martorelli, 1896, p. 279) .
European Lynx. Lynx (Fr.). Luchs (Ger.). Lince (Sp., It.)
LYNX LYNX LYNX (Linnaeus)
[Felis] Lynx Linnaeus, Syst. Nat., ed. 10, vol. 1, p. 43, 1758. (Near Upsala,
Sweden.)
SYNONYM: Felis borealis Thunberg (1798).
FIGS.: Wolf, 1867, ser. 2, pi. 6; Elliot, 1883, pi. 39; Martin, 1910, p. 117, fig. 34;
Colosi, 1933, p. 41.
The Lynx has suffered rather serious depletion of numbers in
its European range, and even total extermination in some of the
countries (Britain, France, Denmark, Switzerland, Italy, Austria,
Hungary). In northern Asia, however, its status remains much
more satisfactory, especially in those areas where the human popu-
lation is still sparse.
The form is heavier than in Felis silvestris; the legs are relatively
longer, the feet more robust, and the tail shorter; upper parts and
sides varying from yellowish brown to brownish gray; back and
sides never thickly spotted; cheeks not conspicuously whiskered;
ears conspicuously tufted at tip (Miller, 1912, p. 472). Head and
body, 800-1,200 mm.; tail, 190-220 mm. (Didier and Rode, 1935,
p. 289).
The range of this Lynx includes the forested portions of Europe
and Asia: north to the tree limit; south to France, northern Italy,
Yugoslavia, Greece, Bulgaria, the Ukraine, central Russia, the Altai
Mountains, northern Mongolia, northern Manchuria, and northern
Korea ; east to Sakhalin and eastern Siberia.
"The European Lynx, Felis lynx, was ... an inhabitant of
Britain in the Pleistocene age, and survived until recent times, and
may even have lingered into the historic period" (Millais, 1904,
p. 168).
266
EXTINCT AND VANISHING MAMMALS
Although the French authors record the Lynx from the Pyrenees
(where it is now probably exterminated), Cabrera suggests (1914,
p. 210) that the animal formerly occurring there was the Spanish
Lynx (Lynx pardellus) . Trouessart in 1884 (p. 229) considered the
European Lynx still present in the Alps, the Jura, and the Pyrenees.
There is a record (the last?) for the Jura in 1834 (Martin, 1910,
FIG. 29. European Lynx (Lynx lynx lynx)
p. 118). Didier and Rode (1935, pp. 290-291) cite records from the
French Alps as late as 1907, 1913, and 1922, but conclude that the
species has probably disappeared from the entire country.
In Germany, for several centuries past, the Lynx has occurred
only as an occasional straggler. From 1773 to 1796 five were shot
in the Thuringian Forest. A few were taken in Upper Silesia at the
beginning of the nineteenth century. Two were shot in the Harz
Mountains in 1817 and 1818. (Blasius, 1857, p. 176.) The species
was exterminated in Pomerania in 1738; in Westphalia in 1745; in
Gotha in 1819; in Bavaria in 1850. In East Prussia several were
taken about 1870, and the animal still occurs frequently on the
eastern boundary. (Krumbiegel, 1930, p. 6.) One was taken in Thur-
ingia in 1843, and one in Wiirttemberg in 1846 (Internationale Ge-
sellschaft zur Erhaltung des Wisents, in Hit., October, 1936). At
ORDER CARNIVORA: CARNIVORES 267
present the law gives it absolute protection (Reichsstelle fiir Natur-
schutz, in Hit., October, 1936) .
In Denmark the species is known only from Stone Age and Bronze
Age remains (Winge, 1908, p. 117) .
In Switzerland it abounded during the seventeenth century, and
numerous captures were made up to the early part of the nineteenth
century. Thereafter it suffered a pronounced decrease. In the
1860's it was still found, but only occasionally, in Grisons, Ticino,
and Valais (one record in 1867). (Fatio, 1869, p. 280.) One was
killed in the Engadine in 1872. The species is now extinct in Switzer-
land. (Federal Forest, Game, and Fish Inspection, Berne, in litt.,
March, 1937.)
The species may be considered extinct in Italy, the last specimens
having been killed in Piedmont in the second half of the last century
(Laboratorio di Zoologia Applicata a Caccia, in litt., September,
1936). De Beaux (1932, p. 9) speaks of the forest of Langhe in the
Maritime Alps, in the province of Cuneo, as its last refuge.
In Austria the Lynx is entirely exterminated. In Carinthia one
was killed in 1848, and another was seen in 1878. In Lower Austria
it was fairly distributed up to the middle of the last century. In the
Tyrol it was very common in the sixteenth century, but during
the next century it decreased decidedly, and at the beginning of the
nineteenth century it was said to be not very rare ; the last one was
killed at Graun in 1873. In Vorarlberg the last one was killed in
1918. (G. Schlesinger, in litt., March, 1937.)
Before World War I the Lynx was found everywhere in the Car-
pathian forests of Hungary (now Czechoslovakia) (J. Schenk, in
litt., November, 1936) . In the higher elevations of the Tatra Range
(Czechoslovakia and Poland) there are a few Lynxes (Maurice,
1927, p. 21). (See also under Poland.)
In the present Hungary the species is not found (J. Schenk, in litt.,
November, 1936) .
During the past century the Lynx was exterminated throughout
Yugoslavia except in the high mountain region of Shar Planina, in
southern Serbia, where four or five specimens are killed every year
(M. Hirtz, in litt., December, 1936).
In Greece it occurs in Epirus, Thessaly, Macedonia, and Thrace.
No decrease has been observed. (Game Department, Greek Ministry
of Agriculture, in litt., October, 1936.)
In Bulgaria "the lynx is about gone, though I saw two fine skins
last week at a fur shop in Varna, on the Black Sea, and one in the
peasant market in Sofia last year." A reserve for the Lynx in the
Pirene Mountains is advocated. (H. W. Shoemaker, in litt., June
30, 1932.) According to the Bulgarian Game Association "Sokol"
(in litt., February, 1937) , the last specimen was killed in 1907. The
268 EXTINCT AND VANISHING MAMMALS
cause of its disappearance was the diminution of the forests (espe-
cially the virgin forests) .
In Rumania the Lynx is an increasingly rarer species. Its dis-
tributional area is the coniferous-forest zone narrower on the
western slope of the eastern Carpathians (Nasaud district), and
broader on the eastern slope (Bukowina) . It is more common in the
southern Carpathians (Bunzenlander Mountains, Fagaras Moun-
tains, etc.). (Calinescu, 1930, p. 366.)
In ancient times, when Poland was covered with large forests, the
Lynx was found everywhere. It still exists in the Carpathians, espe-
cially in the eastern Polish part, and also in the great forests of
northeastern and eastern Poland. The estimated number is about
400. During the hunting season about 25 specimens are killed
annually. Hunting is forbidden from March 1 to December 31. (M.
Siedlecki, 1 in'litt., October, 1936.) In the proposed International
Tatra Park (Poland and Czechoslovakia) the Lynx will be kept
under absolute protection. The chief aim of another proposed Inter-
national Park (Poland, Czechoslovakia, and Rumania) is the crea-
tion of a breeding ground for the Bear, Lynx, Wildcat, Wolf, Stag,
etc. (Benedyct Fulinski, MS., 1933.)
In Lithuania the Lynx is almost exterminated. Since the Great
War about ten specimens have been killed. Hunting is forbidden.
(T. Ivanauskas, in litt, November, 1936.)
In Latvia the species has decreased considerably but is still found
in the large forests. The following numbers have been reported
by the Forest Department: 74 in 1925; 49 in 1932; 59 in 1935; 78
in 1936. (N. von Transehe, in litt., February, 1937.) The animals
are found especially in the northeastern part of the country, where
hunting is restricted; in other parts no protective measures have
been adopted. Twenty to thirty years ago the stock was about
300-400, but for economic reasons, and in order to protect useful
game, the numbers have been reduced intentionally, and they are
now confined to a certain part of the forests, where they are pro-
tected. (Forest Department, Latvia, in litt., March, 1937.)
In Estonia the Lynx is only a visitor, coming in from Russia, and
is given no protection (Zoological Institute, University of Tartu,
in litt., October, 1936) .
i "Michel Siedlecki, Professor of Zoology in the University of Cracow, was
... an enlightened apostle of Fauna Preservation. . . .
"Michel Siedlecki was done to death in one of Germany's most notorious
concentration camps. . . .
"In . . . international relations Siedlecki inspired among his colleagues both
respect and affection. One wonders with what feelings the German colleagues
with whom he collaborated in the International Council for the Exploration
of the Sea, as well as in the International Committee for Bird Preservation,
reflect upon the death, brought about through the calculated brutality of their
Government, of a loyal colleague who was so recently their guest." (Jour. Soc.
Preservation Fauna Empire, n. s., pt. 39, pp. 15-16, 1940.)
ORDER CARNIVORA: CARNIVORES 269
The species was formerly common in the Norwegian forests
north to about latitude 65 30'. A quarter of a century ago Collett
remarked on a great decrease. Since it preys upon the more im-
portant smaller game and on small cattle, it is hunted throughout
the country and is not protected by law. (Hj. Broch, in Hit., Decem-
ber, 1936.) Up to about 1875 it was generally distributed in the
woodlands of Norway. A small stock remains in some places in
Fosen, Namdalen, and South Helgeland. In southern Norway it is
doubtful if there is any resident stock, but now and then some
stragglers may appear. Up to the present, bounties have been paid
on the Lynx, and it is doubtful if its total extermination in Norway
can be prevented. (Director of Forestry, Norwegian Ministry of
Agriculture, in litt., January, 1937.)
About a century ago the Lynx was rather common throughout
Sweden except in the northernmost provinces and in some of the
southernmost. From 1827 to 1839, 3,224 Lynxes were killed an
annual average of 248; in 1844, 250; and in 1845, 273. The annual
average was about 175 in 1856-60; 121 in 1865-69; 105 in 1871-75;
67 in 1876-80; 24 in 1881-85; 35 in 1891-1900; 10 in 1901-05; 17
(all in the four northernmost provinces) in 1906-10; 11 in 1921-25.
In the southern provinces the species was practically exterminated
before 1870; in the middle provinces it was found in diminished
numbers in the 1890's. The increase in numbers killed in the 1890's
and in 1906-10 was due to livelier persecution in the more northern
provinces. 1 In 1926 and 1927, 7 Lynxes were killed. Since then the
species has been protected by law to the extent that it may not be
killed on public domains, and, if killed on private lands, it is never-
theless crown property, so that the profit motive is eliminated. The
crown domains are very extensive, especially in the northern prov-
inces, and there are some state forests, especially in Westerbotten,
where the animals seem to thrive and to increase somewhat. There
are a few in Angermanland and perhaps also in Jemtland, while
stragglers have been traced in some other provinces. (Einar Lonn-
berg, in litt., October, 1936.)
The following information on the Lynx's range and status in
Russia and Siberia is derived from Ognev (1935, pp. 206-214) :
The range extends entirely across Russia from east to west; in the
north it reaches the tree limit on the Kola Peninsula and at other
points near the Arctic coast; in the south it reaches Podolia, the
southern part of the Western and the Moscow Areas, the district
of Penza, and the former Governments of Kazan and Orenburg.
Within this range its numbers vary considerably but are evidently
1 Possibly these increases represented peaks in a periodic fluctuation, such
as is evident in the case of the Canada Lynx; 1895-97 and 1905-06 were peak
periods for the latter species (cf. Seton, 1929, vol. 4, p. 711). Ed.
270 EXTINCT AND VANISHING MAMMALS
much less than formerly. In White Russia 76 animals were taken
in 1924-25, but only 16 in 1925-26. In the Western Area 156 were
killed in 1928-29, 43 in 1929-30, and 47 in 1930-31. According to
Milovanowicz (1925), 50 are taken annually in the Petchora region.
In the Vichegda Basin 3 were taken in 1929, 2 in 1930, and 27
in 1931.
In Siberia the Lynx ranges north approximately to the tree limit
and south to northern Russian Turkestan (Irtish River and Zaisan
Nor) and to the Mongolian and Manchurian boundaries, apparently
avoiding the Arctic tundra on the one hand and the steppes of Rus-
sian Turkestan on the other. (In far eastern Siberia, beyond the
Verkhoyansk Mountains, the typical subspecies seems to be replaced
by Lynx lynx wrangeli Ognev.)
According to W. G. Heptner (in litt., December, 1936) , the Lynx
is quite common in Siberia but rarer in Russia. Hunting is allowed
the whole year, but in certain parts of Russia and in western Siberia,
where the numbers of the animal are small, hunting is limited to
certain open seasons.
The species occurs in the mountains of northern Mongolia, from
the Altai Range eastward (Ognev, 1935, p. 214) . A specimen from
15 miles northeast of Urga, Mongolia, is recorded by G. M. Allen
(1929, p. 14) under the name of L. I. isabellina. The species also
occurs in the forests of Manchuria; it is rare in northern Kirin
(Sowerby, 1923, p. 37) and in the southern part of the Little Khin-
gan Mountains (Ognev, 1935, p. 213). It is generally considered
rare in Sakhalin (Kuroda, 1928, p. 226; Miyoshi, in Skottsberg,
1934, p. 411).
Economics. The Lynx attacks game as large as the Red Deer
and the Roe Deer, and it has been known to slaughter 30 sheep in a
single night. Generally it feeds on the smaller game hares, mar-
mots, small rodents, and birds of all kinds. On occasion it does not
fear to attack man himself. (Trouessart, 1884, p. 230.)
In Sweden, "when the lynxes were numerous, they sometimes killed
sheep and perhaps also reindeer. They are therefore like all other
carnivorous animals especially hated by the Lapps. The lynxes
were also destructive to the hares, when they were common. The
reason why they were so much hunted was, however, chiefly because
it was regarded as a good sport, and the value of the skin was also
attractive." The species is now protected in Sweden, and the govern-
ment pays for the damages committed, "if they are not due to
carelessness of the owners of the domestic animals killed." (Einar
Lonnberg, in litt., October, 1936, and January 18, 1933.)
"The Norwegian peasants believe that if a person wears a neck-
lace made of the fore-claws of a lynx it will preserve him from
spasms and the cramp. It would appear that the Russians entertain
ORDER CARNIVORA: CARNIVORES 271
a somewhat similar superstition, for when they sell lynx skins to the
Chinese, they charge a much higher price for them if the fore-claws
are included." (Bowden, 1869, p. 14.)
In the Polish forests "the lynx and the wolf alone are capable
of killing the elk" (Korsak, 1934, pp. 78-79).
"The Russian naturalists Von Schrenk and Radde inform us that
the natives of Amoorland esteem the flesh of this animal as a great
delicacy, and that the furs which are obtained by the hunters in
this part of Asia mostly pass into Chinese hands, being much trea-
sured by the high officials of the Celestial Empire" (Sclater, in Wolf,
1867, text to pi. 6).
Tibetan Lynx
LYNX LYNX ISABELLINUS (Blyth)
Felis isabellina Blyth, Jour. Asiatic Soc. Bengal, vol. 16, p. 1178, 1847. ("Tibet.")
FIGS.: China Jour., vol. 23, no. 3, pi. facing p. 172, 1935, and vol. 25, no. 5,
pi. facing p. 288, 1936; Schafer, 1937, pi. facing p. 177.
The Tibetan Lynx seems to be a moderately rare animal, and
while it is protected to some extent by the remoteness of its haunts,
its fur is in considerable demand.
It differs from the European Lynx in its pale sandy gray or
isabelline coloring, and in the relative shortness of the hair on the
toes. Head and body, 837 mm. ; tail, 196 mm. ; weight, about 60 Ib.
(Blanford, 1888-91, p. 90.)
"This race inhabits the plateau of Eastern and Western Tibet,
and certainly extends into Baltistan; but its exact geographical
limits are impossible to define ....
"Throughout its habitat, so far as accounts go, the Tibetan lynx
is a rare animal, seldom seen, and still more rarely shot. . . . The
Tibetan hares and blue pigeons form the chief prey of the lynx in
Ladak, although it also levies toll on the smaller domesticated ani-
mals of the Tatars." (Lydekker, 1900, pp. 326-327.)
"This animal is rarely encountered and consequently the exact
limits of its habitat are somewhat conjectural, but I fancy that it is
almost identical with that of Ammon. . . . They are savage ani-
mals and do not hesitate to attack sheep and goats, sometimes
working considerable havoc." (Burrard, 1925?, p. 241.)
"Lynx skins . . . are brought in from the Thibetan regions to
the north and west, to Sungpan [Szechwan] , where they find a ready
market among the wealthy Chinese. . . . They sell in Sungpan
for 5 to 7 taels each." (Wilson, 1913, vol. 2, p. 181.)
"The Isabelline Lynx ... is fairly common in the mountainous
regions along the Chinese-Tibetan border. . . . The lynx supplies
the fur market of this country [China] with one of its best furs, the
272 EXTINCT AND VANISHING MAMMALS
coat of this animal being long, thick and soft." (Sowerby, 1936,
pi. facing p. 288.)
Ognev (1935, pp. 215, 231-232) reports it from Kansu, Koko Nor,
the Zaidam region, the Nan-Shan, Altyn-Tagh, Kwen-Lun, Tian-
Shan, Borokhoro, and Bogdo-Ola ranges, Kashgar, the Tarim Basin,
and Lob Nor.
The systematic status of the Lynxes inhabiting various mountain-
ous areas from northwestern India and southern Russian Turkestan
to northern Persia seems to be in doubt; I am unable to determine
what subspecific name or names should be applied to them. This
vast region lies directly between the ranges of the Tibetan Lynx
and the Caucasian Lynx.
The animal of Gilgit, in the valley of the Indus, has a more
rufous coloring than that of Tibet (Lydekker, 1900, p. 326) .
According to Ognev (1935, pp. 214-215), the Lynx is found in
various localities of Russian Turkestan, including the Pamirs, the
western Tian-Shan, Semiretchie, the Chu River, the Kara Tau, the
Talassk Alatau, the Samarkand region, and the Kopet-Dagh (where
it is rare). It occurs in northern Afghanistan and doubtless in the
Persian provinces of Gilian, Mazanderan, and Astrabad; possibly
also in the mountains of Khorassan (lat. 37 N.).
W. G. Heptner states (in litt., December, 1936) that the Lynx
occurs in small numbers in the mountains of Turkestan. Hunting
is limited to certain seasons in the mountains of Uzbekistan, and is
forbidden on the Kopet-Dagh.
Barbary Lynx. Lynx caracal (Fr.)
CARACAL CARACAL ALGIRUS (Wagner)
Felis Caracal . . . Var. algira Wagner, Reisen Regentschaft Algier, vol. 3,
p. 76, atlas, pi. 4, 1841. (Vicinity of Algiers (op. tit., p. 62).)
FIGS.: Buffon, Hist. Nat., vol. 9, pi. 24, 1761; M. Wagner, 1841, atlas, pi. 4;
Loche, 1867, pi. 2.
This Caracal is evidently becoming increasingly scarce as the
years roll by.
General color nearly uniform, between cinnamon-orange and
reddish cinnamon; paler about the eyes, on the lips, and on the
lower parts; a blackish spot on each side of the mouth; ears ex-
ternally black, sometimes mixed with white hairs, terminal tuft
black. Head and body, 717 mm.; tail, 284 mm. (Cabrera, 1932,
pp. 171-172.)
The species as a whole is "widely distributed in suitable localities
from South Africa to Egypt and Morocco, and from Palestine to
India" (Flower, 1929, p. 83). Only the North African subspecies
ORDER CARNIVORA: CARNIVORES 273
(algirus) calls for attention here. It ranges from the Gulf of Gabes
to the Atlantic coast, and south to Senegal. In Algeria and Tunisia
it reaches the Mediterranean coast. Apparently it is not represented
in Tripolitania. (Cabrera, 1932, pp. 172-173.)
Tunisia. It is rare in the north but a little more common in the
center and south (region of Feriana, Djebel-Selloum, Djebel-Bou-
Hedma). It can be shot rather easily in its usual habitat among
alfa grass, and it could be successfully chased with hounds, for its
gait is not very rapid. (Lavauden, 1932, p. 7.)
FIG. 30. Barbary Lynx (Caracal caracal algirus). From specimen in
Philadelphia Zoo.
Algeria. The Barbary Lynx does not occur commonly in the
vicinity of Algiers, whence occasionally specimens are brought to
market (M. Wagner, 1841, vol. 3, p. 62).
Loche (1867, p. 41) reports for the period 1840-42 that it ranges
throughout Algeria, where it is rather numerous. Specimens are
recorded from Birkadem, Arba, and Djelfa. It is also met with near
Coleah, Lac Halloula, and elsewhere.
Known in Barbary from the first explorations, it has been recorded
by Shaw, Poiret, and others. Two specimens were taken in the
vicinity of Laghouat, and one of the animals was seen between
Haidra and Tebessa, Algeria. Without being very abundant, the
species seems rather widespread, and skins are frequently seen in
the saddlers' shops of Algiers and Constantine. (Lataste, 1885,
p. 225.)
10
274 EXTINCT AND VANISHING MAMMALS
Heim de Balsac (1936, p. 179) records it from the valley of the
Saura, near Beni-Abbes.
Morocco. "The Lynx ... is found in wooded districts, and is
sometimes brought alive to Mogador" (Leared, 1876, p. 304). It
seems to be lacking in Yebala, but it undoubtedly exists more to
the south, and it is reported as still living in the interior of the
Rif (Cabrera, 1932, pp. 172-173) . In the Zaian district it is found
from time to time, and its skin is frequently seen in the market. One
was living in captivity as late as 1931. (Carpentier, 1932, p. 17.)
Specimens are recorded from Gara de Debdou, Matarka, Oued
Charef , region of Berguent (Laurent, 1935, p. 349) .
Gambia. "This splendid animal is to be seen some 150 miles up
the [Gambia] river; being swift and cunning, very few are trapped
or shot" (E. Johnson, 1937, p. 63). The local form does not seem
to have been subspecifically determined.
The Cheetahs (genus Acinonyx)
These animals are also known as Hunting Leopards. Additional
names in various Continental languages are: Guepard (Fr.), Gep-
pard (Ger.), Ghepardo (It.), and Onza (Span.).
The common species of Cheetah (Acinonyx jubatus) , which has
been divided by various authors into approximately a dozen differ-
ent forms, has become rather rare over a large part of its enormous
range in Africa and Asia, while remaining moderately common in
some areas. Another species, the King Cheetah (A. rex), has been
described from a restricted area in Southern Rhodesia. For the
sake of completeness, all forms that are more or less recognizable
will be treated in the following accounts. In scarcely any case can
the distributional limits of the subspecies be stated precisely, owing
to the incompleteness of our present knowledge.
The Cheetahs are distinguished from all other members of the
cat family (Felidae) by the absence of claw-sheaths (Pocock, 1916,
p. 426). In size and form they suggest a long-legged and slender-
bodied Leopard; but their markings are solid spots instead of ro-
settes, as in the Leopard. The hair of the neck is elongated to form
a slight mane. (Lydekker, 1900, p. 328.)
The geographical distribution of the Cheetahs as a genus is very
similar to that of the Lion. In Africa it includes chiefly the arid
or semiarid areas of the South, East, and North in fact, most of
the continent outside of the rain forests of the West African sub-
region and the humid, forested areas of southeastern Africa. In
Asia, likewise, it includes more or less arid areas from India and
Russian Turkestan to Syria, Palestine, and Arabia.
ORDER CARNIVORA: CARNIVORES 275
Economics and conservation. "The Cheetah has been for cen-
turies the playing thing of princes, Asiatic, African and European.
When taken young it is easily tamed and trained to show its won-
derful speed. A couple of cheetahs were sent as a present to Ana-
stasius, Emperor of the East, at Constantinople in A. D. 439 from
whence the sport reached Italy and obtained an enormous vogue
among the wealthy, extravagant grandees of the great City States.
The Turks, Moors and Persians alike used cheetahs in hunting. In
India the Cheetah is usually taken blind-folded in a cart to the
scene of the hunt. In the proximity of a herd of antelope it is un-
hooded and slipped from its leash: a short crouching stalk and a
few bounds of great length and rapidity and the hunt is over
the quarry has escaped or the Cheetah holds it in a strangle-hold
by the throat, till the keeper comes up and having cut the captive's
throat rewards the captor with a drink of warm blood collected in
its accustomed feeding bowl." (Anonymous, 1935, pp. 148-149.)
(For an interesting account of hunting with Cheetahs in the
Middle Ages, see Yule's Marco Polo, ed. 3, vol. 1, pp. 397-398,
1903.)
"So far as I have heard, . . . this animal has not been known to
breed in captivity" (Blanford, 1888, p. 93) .
Probably the disappearance of the Cheetah in the Cape Province
and its decrease in North Africa and in Asia are closely linked with
the general decline in those regions of the various antelopes which
constitute its principal prey. It also attacks calves, sheep, and goats
to some extent, and thereby comes into conflict with man. However,
"there are very few recorded cases of cheetah attacking human be-
ings or taking the offensive, even when wounded or in defence of
their cubs" (Shortridge, 1934, p. 107) . Like all of the larger carni-
vores, it is evidently shot "on general principles." According to
Shortridge (1934, p. 107), it retreats more rapidly than the Leopard
before European settlement.
In Northern Rhodesia "Cheetah and other animals take their toll
of the vast numbers of Black Lechwe, but these natural enemies do
more good than harm" (David Ross, in litt., May 3, 1933) .
Fortunately the Cheetah is reported present in a considerable
number of the African game reserves: White Nile Reserve and one
near the Ethiopian border in the Sudan; reserve between the Gash
and Setit Rivers in Eritrea; Northern and Southern Reserves in
Kenya; Katavi, Mtandu, Lake Natron, Ngorongoro, Northern Rail-
way, Saba, Selous, and Serengeti Reserves in Tanganyika; Matupo
Reserve in Mozambique; Kruger National Park in Transvaal;
Bechuanaland Reserve in Bechuanaland Protectorate; and Namu-
toni Reserve in South- West Africa. Such reserves doubtless afford
276 EXTINCT AND VANISHING MAMMALS
the species its best chance of survival in countries that continue to
be opened up to settlement and cultivation.
Cheetahs make charming pets, and may be led about on a leash.
South African Cheetah
ACINONYX JUBATUS JUBATus (Schreber)
Felis jubata Schreber, Saugethiere, vol. 3, pi. 105, 1776; p. 392, 1777. (Cape
of Good Hope, South Africa.)
SYNONYMS: Felis lanea P. L. Sclater (1877); lAcinonyx guttatus obergi
Hilzheimer (1913).
FIGS.: Proc. Zool. Soc. London 1877, pi. 35 (incipient albinism); Hilzheimer,
1913, p. 289, fig. 2 (A. j. obergi) 1
The South African Cheetah has virtually disappeared from the
Cape Province. Elsewhere in its huge range it has managed to sur-
vive in varying numbers, but in general it is obliged to retreat before
settlement.
The general color is ochraceous-yellow, covered with round black
spots; below almost white, with longer hair and indistinct spots; a
black line from the anterior angle of the eye to the edge of the lip ;
ear with a black patch posteriorly and a tawny tip; chin white,
unspotted; chest whitish, spotted; spots on the tail tending to form
6-8 imperfect rings toward the tip. (W. L. Sclater, 1900, p. 47.)
Total length, about 6 feet 8 inches; tail, about 2 feet 6 inches;
shoulder height, about 2 feet 8 inches.
All Cheetahs (except A. rex Pocock) occurring from the Cape
Province north to Angola, Belgian Congo, Northern Rhodesia, and
Nyasaland will be included provisionally under the present sub-
species. This range corresponds roughly to the Southwest Arid
District, the Southeast Veld District, and the Rhodesian Highland
District of Chapin (1932, p. 90), or to the Southeast Veldt District,
the Kalahari and the Damara Arid Districts, and the Rhodesian Sa-
vanna District of Bowen (1933, pp. 256, 259-260) .
"In the [Cape] Colony it is found sparingly in the western and
midland districts : north of the Orange River it is common in German
territory [South-West Africa], the Kalahari and Bechuanaland, and
exists in Rhodesia, the Transvaal, Zululand and Natal, though now
very rare in the latter Colony, and found only in the Drakensberg
range" (W. L. Sclater, 1900, p. 48) .
In South-West Africa the Cheetah is widely distributed. It is
considered quite plentiful in the eastern sand-veld region, scarce
in the Kaokoveld, sparsely distributed in the Caprivi, and not un-
common in the Namutoni Game Reserve. It is recorded from various
i The following additional figures of Cheetahs are of undetermined subspecific
identity: Wolf, 1861, pi. 13; Lavauden, 1924, p. 10, fig.; Leplae, 1925, p. 46,
fig.; Malbrant, 1936, pi. 1, upper fig.
ORDER CARNIVORA: CARNIVORES 277
parts of Omaheke. It is apparently scarce in the vicinity of the
Orange River, the western and southwestern parts of Great Nama-
qualand, and the highlands of western Damaraland. Its numbers
increase in the sand-plain country adjoining Bechuanaland. Con-
siderable numbers of skins are brought into Windhoek and Keet-
manshoep annually by natives from this region and from Bechuana-
land. The species also occurs in small numbers in southern and
southeastern Angola. (Shortridge, 1934, p. 105.) This author adds
(pp. 105-107) :
The Cheetah has almost if not completely disappeared from the Cape
Province, the Orange Free State, Natal, and the Southern Transvaal, but
may still be met with in some of the more sparsely populated districts of the
Northern Transvaal, Zululand, Swaziland, and probably the inland portions
of Portuguese East Africa. It is widely distributed in Bechuanaland and
still comparatively plentiful in the central and northern portions of that
territory. . . .
The Cheetah is retreating rapidly before settlement in Southern Africa, and
it is doubtful if there are any to be found to-day south of the Vaal River. . . .
Cheetah prey mostly upon medium-sized antelope, from steinbok and duiker
up to the size of impala, springbok, reedbuck, and even cow kudu. . . .
When opportunity offers they kill sheep, goats, and ostriches, which last
are driven into wire fences and cornered.
In the Transvaal the species is considered inimical to man, and
not a game animal. Thus it is not given any protection. (Austin
Roberts, in Hit., November, 1936.)
In the Kruger National Park, Transvaal, "the status of the species
remains fairly constant; they were never very numerous, and no
noticeable increase or decrease is reported" (Game Warden's Annual
Report, 1925?).
In Southern Rhodesia Cheetahs are sparingly distributed over
the greater part of the country, but soon retire from inhabited
areas. They seldom raid domestic stock and consequently do not
often fall a victim to traps and poison. There is no legal protection,
but in the recognized game reserves and also in the forest reserves all
animals are rigidly protected. (Game Warden, Wankie Game Re-
serve, in Hit., March, 1937.)
In Northern Rhodesia this "widely distributed species ... is
absent from the regions of interminable woodland. Usually occurs
sparingly, though inclined to be locally plentiful." It "is most nu-
merous in the neighbourhood of open expanses such as the Kafue
flats, the Batonga and Batoka plateaux, the neighbourhood of Bang-
weulu, the Chambeshi flats and other similar localities. It appears
to be absent from a great part of the Luangwa Valley. Family
parties up to five are frequently reported, and as many as seven have
been seen together. The cheetah is a very disturbing factor in locali-
ties where it occurs side-by-side with domestic stock, and is apt to be
278 EXTINCT AND VANISHING MAMMALS
particularly destructive where sheep and calves are concerned."
(Pitman, 1934, p. 12.) Skins have been obtained in the Mumbwa,
Namwala, Broken Hill, Mpika, Chinsali, and Petauke Districts
(op. cit., p. 159).
In Nyasaland "the Cheetah has so far only been found in the
Central Province of Angoniland. Even there it is seldom seen, but
may be more numerous than is believed at present, as it is nearly
always confused with the Leopard by the natives." (Wood, in May-
don, 1932, p. 316.)
In Portuguese East Africa "Cheetahs ... are not very numerous
in the Zambezi valley, occurring perhaps most plentifully between
Muterara and the Lupata Gorge, where reed buck and other small
antelopes are common. I have also seen them in the Mlanje district
. . . , in the Barue to the south of Tete, and in the open country
south of the Shupanga Forest." (Maugham, 1914, p. 195.)
In the Belgian Congo, 20 years ago, the species was comparatively
abundant over the southern part of the colony, from Kwango to
Tanganyika. It probably ranged to the northern extremity of Lake
Tanganyika, wherever grassy stretches, inhabited by small rumi-
nants, assured it of favorable conditions for existence. The range
seems to have remained practically the same, except in southern
Katanga and Lomami, where stock-raising has led to the extermina-
tion of Cheetahs. It is well to note that these stock farms have
been established where Cheetahs were particularly numerous. Fur-
thermore, these animals suffer greatly from hunting with encircling
fires, and the natives persecute them everywhere to satisfy the
demand for skins on the part of the European population. The
Cheetah should be put on the protected list. That would suffice, if
the customary destructive hunting by the natives could be effectively
stopped, to assure the recuperation of the species outside the zones
of stock-raising. (A. J. Jobaert, in litt., November 10, 1936.)
East African Cheetah
ACINONYX JUBATUS NGORONGORENSis Hilzheimer
A[cinonyx\ g[uttatus] ngorongorensis Hilzheimer, Sitz.-ber. Ges. naturf.
Freunde Berlin 1913, no. 5, p. 290, figs. 3-4, 1913. (Based upon a living
specimen in the Leipzig Zoological Garden, said to have come from
"Ngorongoro," south of Lake Natron, Tanganyika Territory.)
FIGS.: Hilzheimer, 1913, pp. 290-291, figs. 3-4; Roosevelt and Heller, 1914,
pi. facing p. 244 (raineyi) ; Hollister, 1918, pi. 5, lower fig. (raineyi) ;
Zammarano, 1930, p. 152, fig. (ngorongorensis?).
Two other forms were subsequently described from East Africa:
A. j. velox Heller, Smithsonian Misc. Coll., vol. 61, no. 19, p. 7, 1913
("Loita Plains, British East Africa"), and A. j. raineyi Heller,
Smithsonian Misc. Coll., vol. 61, no. 19, p. 9, 1913 ("Ulu, Kapiti
ORDER CARNIVORA: CARNIVORES 279
Plains, British East Africa"). Hollister (1918, p. 151) seems to be
somewhat doubtful as to the distinctness of these three East African
forms from each other, and De Beaux (1927, pp. 3-4) does not
recognize velox or raineyi. The Cheetahs of East Africa, from
Tanganyika north to Ethiopia and Eritrea, will be treated here
as a unit. This range corresponds roughly to the Somali Arid District
and the East African Highland District of Chapin (1932, p. 90), or
to the Somali Arid District, the North Kenya Savanna District,
and the East African Highland District of Bowen (1933, pp. 256,
258,260).
The ground color of the living type specimen of ngorongorensis
was isabella yellow-brown; under parts very light isabella, entirely
without white; spotting of the lower neck very pronounced; cheeks
grayish, heavily spotted; back of the ear yellow, with a slender
black basal stripe; chin and lips white; chest and belly unspotted;
outer side of the limbs heavily spotted as far as the toes; tail with
three complete rings, tip yellowish white (Hilzheimer, 1913, p. 289) .
The coloration of this captive specimen may not have been typical.
The following might serve as a composite characterization of velox
and raineyi: ground color ochraceous to light pinkish buff; spots
large, blackish; snout ochraceous to ochraceous-buff ; a black tear
stripe from eye to mouth; back of ears black basally, tip and inner
surface buff or pinkish buff; chin and upper throat white to cream-
buff ; belly cream to cream-buff, with elongate spots ; hind feet more
or less distinctly spotted; terminal part of tail ringed with black,
tip whitish (Heller, 1913, pp. 8-10). Head and body, 1,120-1,300
mm.; tail, 720-800 mm. '(Hollister, 1918, p. 154).
In East Africa and in South-West Africa the Cheetah seems to
have survived in more satisfactory numbers than elsewhere in its
wide range.
It occurs throughout Tanganyika Territory in varying numbers.
There is no danger of extinction at present. In five provinces out of
eight, only two Cheetahs may be killed on a Full Licence, and only
one on a Minor Licence. (Game Preservation Department, Tangan-
yika Territory, in litt., December, 1936.) It is "quite numerous in
Masailand" (Browne, in Maydon, 1932, p. 312).
In Kenya the Cheetah is fairly common and generally distributed,
specimens being recorded from the Loita, Kapiti, and Athi Plains,
Laikipia and Uasin Gishu Plateaus, the flanks of Kilimanjaro, and
Upper Tana River (Roosevelt and Heller, 1914, pp. 244-249) . There
are fair numbers in the Southern and Northern Game Reserves (Per-
cival, 1923, pp. 69-71). There is a decrease in the Native Reserves
and in the European settled areas; otherwise it is fairly common,
and it is protected (Game Warden, Kenya, in litt, November, 1936) .
280 EXTINCT AND VANISHING MAMMALS
The Cheetah is sparingly distributed throughout northern and
eastern Uganda. Not long ago it had a much more extensive distri-
bution, occurring throughout the savanna regions. Its disappearance
from many localities is due to the extension of settlement and culti-
vation. The Cheetah is of sentimental importance and also of
considerable economic value, being sought after for hunting ante-
lopes in India. Specimens trained by their parents in the field have
a local value of 20 to 30. Only one specimen is allowed on a
Full Game Licence. (Game Warden, Uganda, in litt., December,
1936.)
De Beaux (1927, p. 4) records the species from Italian Somaliland,
Ethiopia, and Eritrea. In the southern plain of Eritrea, especially
between Barca and Gash, the animal is rather frequent (Zamma-
rano, 1930, p. 61).
In British Somaliland "the cheetah is commonest in the thick
bush country on the edge of the Haud, although it is to be found
both on Guban and Ogo-Guban" (Drake-Brockman, 1910, p. 22).
Sudan Cheetah
ACINONYX JUBATUS SOEMMERINGII (Fltzinger)
Cynailurus Soemmeringii "Riippell" Fitzinger, Sitz.-ber. math.-nat. Cl. Akad.
Wiss. [Wien], vol. 17, Heft 2, p. 245, 1855. (Based upon a living specimen
from the Kababish Steppes in the south of the Bayuda Desert, Kordofan.)
Roosevelt and Heller (1914, p. 249) give the range as "lowlands
of the Nile Valley, from the Albert Nyanza northward to Kordofan
and westward to Lake Chad and northern Nigeria." No information
is at hand as to the exact northern or western limits of this sub-
species, where it should presumably intergrade with A. j. hecki.
Its range lies in the eastern portions of the Sudanese Arid District
and the Sudanese Savanna District of Chapin (1932, p. 90) and of
Bowen (1933, pp. 256, 258).
The ground color above is ochraceous or pinkish buff; spots not
exceeding half an inch in diameter and widely separated ; hind feet
unspotted. However, according to Malbrant (1936, p. 137 and pi. 1,
upper fig.), the hind legs are nearly always spotted in Cheetahs of
the Chad region.
Roosevelt and Heller (1914, p. 250) record specimens from El
Dueim on the White Nile and from Lake Chad. "It is a rare animal
in the Nile district and is seldom secured by sportsmen. . . . Heller
saw a pair near Gondokoro."
"Cheetah, although by no means common in the Sudan, are widely
distributed throughout the country. They are even reported to
exist as far north as Jebel Tegaru in the north-west corner of the
Province of Kordofan." (Brocklehurst, 1931, p. 32.)
ORDER CARNIVORA: CARNIVORES 281
Butler (in Maydon, 1932, p. 139) refers to this Cheetah in the
Blue Nile district as "everywhere a much scarcer beast than the
Leopard, and rarer in the Eastern Sudan than it is in Kordofan.
Indeed, on this side of the country I only met it twice, both times
on the Setit."
"Heuglin gives the following locality for the Chitah: Southern
Takah and Eastern Sudan not north of 19 N. ... Ruppell . . .
mentions Felis guttata as one of the animals hunted ... in the
western deserts of the Dongola district." (Anderson and de Winton,
1902, p. 185.)
In the Ubangi-Shari district of French Equatorial Africa a few
Cheetahs may occur in the extreme north (Birao), but it is not
certain. They are not threatened for the moment." (L. Blancou,
in litt., December, 1936.)
In the French Cameroons the species is found in the thorn-bush
country, but is very rare. It does not have any special legal pro-
tection. (Ministry of Colonies, Paris, in litt., November 7, 1936.)
In writing of the Chad region, Malbrant says (1936, pp. 137-138)
that the Cheetah is found in the whole of the Sahelian region of
central Africa and in the somewhat forested steppes, its southern
limit being at about lat. 10 N. It lives sometimes solitarily, but
more often in bands of two to four individuals. The natives of Chad
do not utilize it for the chase.
It is found in French Sudan, the Niger Territory, Borku and
Ennedi, and the desert part of the Chad Territory (General Gov-
ernment of French West Africa, in litt., November, 1936) .
In Nigeria "it is pleasant to be able to report that cheetah are
not nearly as rare as was thought. There are fair numbers in several
Provinces and they extend nearly as far south as the Benue River."
("Observer," 1934, p. 54.)
Senegal Cheetah ; North African Cheetah
ACINONYX JUBATUS HECKi Hilzheimer
Acinonyx hecki Hilzheimer, Sitz.-ber. Ges. naturf. Freunde Berlin 1913, p. 288,
fig. 1, 1913. (Based upon a living specimen in the Berlin Zoological Garden,
said to have come from Senegal.)
Fios.: Geoffroy and Cuvier, Hist. Nat. Mamm., vol. 3, pi. 145, 1824; Hilz-
heimer, 1913, p. 287, fig. 1.
The name A. j. hecki, although based upon a Senegal specimen,
may be provisionally applied to the Cheetah occurring over the
greater part of Palaearctic Africa (Mauretania, Morocco, Algeria,
Tunisia, Cirenaica, and northwestern Egypt). It is a rare form.
It is described as a small, dainty animal, with a ground color of
pale reddish ochraceous on the back and sides; spots mostly black,
282 EXTINCT AND VANISHING MAMMALS
but brownish on the cheeks, hind feet, and part of the forefeet;
under parts white, unspotted except on the lower neck; tail with
four complete rings and a white tip (Hilzheimer, 1913, pp. 287-
288). Head and body, 1,150 mm.; tail, 650 mm. (Cabrera, 1932,
p. 192).
In Senegal the Cheetah is met with as far south as Podor and
even near St. Louis (Cligny, 1900, p. 289).
In Morocco it exists only in the Saharan district, south of the
Grand Atlas and the Anti-Atlas. Strohl (1923) refers to the capture
of a dozen specimens in the vicinity of Zenaga [region of Figuig?].
According to native report, it is well known, though not very com-
mon, in the Wadi Draa. Thence it extends across Mauretania to
Senegal. (Cabrera, 1932, p. 192.) Laurent (1935, p. 350) records
skins from Tamlelt, Morocco.
In North Africa the Cheetah is extremely rare, but is still found
regularly on the Oran-Moroccan High Plateaus. It is also said to
be distributed here and there in the entire Sahara. (Heim de Balsac,
1936, pp. 99, 179.)
In Tunisia this very rare animal exists only in the extreme south,
in the Grand Erg. Sometimes solitary individuals range toward
the north; thus some Cheetahs were killed at Fedjej and at El-
Hamma in 1908 and 1913. The species is hunted by its tracks,
which are easy to follow on the sand of the dunes. The natives of
southern Tunisia do not utilize it for hunting, as the Afghans, Arabs,
and Indians do with the Asiatic Cheetah. (Lavauden, 1932, pp. 7-8.)
The cause of depletion in Tunisia is the progress of civilization;
there are no special protective measures. The animal is found ac-
companying herds of Addax and Loder's Gazelles. (Conservator of
Forests, Tunis, in Hit., September, 1936.)
We lack precise information on the range of the Cheetah in the
Libyan hinterland. Some are found in the southern steppe region of
Cirenaica. A specimen was killed recently at Bir Scegga, between
Tobruk and Jarabub. At the time of the Pharaohs the animal was
employed in the hunting of antelopes. Its skin has slight commercial
value. (Zammarano, 1930, pp. 13-15.)
In Egypt the species "is very rare, and found only in the country
to the west of Alexandria. In 1909 Col. H. C. B. Hopkinson . . .
saw the tracks of two Chitas that had been stalking gazelle in the
Mariut district about 40 miles west of Alexandria." In 1910 "a
Bedawin shot a Chita about 5 miles north-east of Moghara, Mariut
district .... A few other specimens were shot later." In 1927
"three live Chita cubs from south-west of Sollum had been received
recently at the Giza Zoological Gardens." (Flower, 1932, p. 392.)
The present range of this vanishing species in Egypt is restricted
to the Western Desert. The cause of depletion is injudicious hunting.
ORDER CARNIVORA: CARNIVORES . 283
The skins are sold and the meat is used for food. (Ministry of
Agriculture and Zoological Garden, Cairo, in litt., January, 1937.)
Indian Cheetah; Indian Hunting Leopard
ACINONYX JUBATUS vENATicus (Hamilton Smith)
F[elis] Venatica Hamilton Smith, in Griffith's Cuvier's Anim. Kingdom,
vol. 5, p. 166, 1827. ("India.")
FIGS.: Jour. Bombay Nat. Hist. Soc., vol. 37, no. 4, suppl., pi. 46, 1935 (venati-
cust); Bodenheimer, 1935, pi. 9 (venations?).
The Cheetah is nearly extinct in India and has become very rare
in southwestern Asia generally.
The general color is pale brownish yellow to bright rufous-fawn
above and on the sides; almost everywhere with small round black
spots ; chin and throat buffy white, unspotted ; a black line from the
anterior corner of the eye to the upper lip, and another less marked
(or a row of spots in some specimens) from the posterior corner of
the eye to below the ear; ear black outside, base and margins tawny;
spots on tail passing toward the end into imperfect rings. Head and
body, about 4.5 feet; tail, 2.5 feet. (Blanford, 1888, p. 91.)
The Cheetahs ranging from Baluchistan, Persia, and Iraq to Syria,
Palestine, and Arabia are here included provisionally with the
Indian form (A. j. venaticus).
In India the Cheetah is all but extinct in the wild state. It once ranged
from the confines of Bengal through the plains of the United Provinces,
the Punjab and Rajputana, through Central India and the Deccan. ... A
Cheetah was killed in 1918 and another in 1919 in the Mirzapur District of
the United Provinces. Five Cheetahs are recorded as having been obtained
in this Province during the previous twenty-five years. In the Central
Provinces, the Cheetah appears to have been not uncommon at one time in
the Berars. Three were shot in the Melghat Forest area in 1890 and one
in 1894 and one at Wano in 1895. Rumours of their existence in parts of
Berar, the Seoni Plateau and Saugor still persist. They were apparently once
common around Hyderabad, Deccan. The only part of the Bombay Presi-
dency where Cheetahs were known to occur recently is the tract of rugged
country known as the Tanga in the centre of the province of Kathiawar. In
1884 it was estimated that there were not more than twenty of these animals
in this area. A female and four cubs were shot at Rajkot in 1894. (Anonymous,
1935, p. 147.)
"In the case of India, the cheetah appears to be verging on ex-
tinction, if not already extinct, as a wild animal. At all events the
Mammal Survey of India . . . does not seem to have secured a
single specimen; and ... it seems that Indian cheetahs are now
practically unobtainable, and that those used for the chase are
imported from Africa." (Pocock, 1927, pp. 18-19.)
"I have heard that Princes and others who want cheetahs for
hunting purposes now get them from Hyderabad. But the officer
284
EXTINCT AND VANISHING MAMMALS
in charge of the Gwalior shikar department . . . said that 50 or
60 survived in the state. They are found in Indore also." (Edward
Thompson, London Times, August 19, 1932?) In Hyderabad State
"there still remain a few cheetahs" (Salim AH, in Anonymous, 1935,
p. 231). In Mysore there are several old records (Morris, 1935, p.
386), but the animal is "probably now extinct" there (Phythian-
Adams, in Anonymous, 1935, p. 241). Fears are entertained as to
FIG. 31. Indian Cheetah (Acinonyx jubatus venaticus)
its survival in the Madras Presidency (M. F. Budge, in litt., Novem-
ber 12, 1936) . Shooting is completely prohibited (Bombay Natural
History Society, in litt., December, 1936) .
"The hunting leopard is certainly found in Persia, but I am unable
to give any particulars as to its distribution. According to Eichwald
it does not extend into the countries west of the Caspian, though
found to the eastward. De Filippi says that it is found in Mazan-
daran." (Blanford, 1876, p. 35.) To this statement 0. St. John adds
that the species "is not used at present for sporting purposes in
Persia." [The form of northern Persia may be A. j. raddei.]
"As regards its survival in Persia, Arabia and Palestine, I have
no information beyond the inference to be drawn from the infre-
quency with which it is mentioned" (Pocock, 1927, p. 18) .
"In South-Western Asia its range is believed to reach from the
ORDER CARNIVORA: CARNIVORES 285
frontiers of Sind through parts of Afghanistan, Baluchistan and
Persia and Mesopotamia to Syria and Palestine .... To what
extent it survives in these Asiatic countries is not known." (Anony-
mous, 1935, p. 147.)
In 1925 a cub was secured at Jumaimah, Muntafq, Iraq, and in
1928 two cubs were taken near Busiya on the Shamiyah Desert.
None of the local Arabs had seen a Cheetah before. (Corkill, 1929,
pp. 700-702.)
Danford and Alston (1880, pp. 52-53) report on the Cheetah in
Syria as follows: "A skin of the Cheetah was presented to Danford
at Biledjik, on the Euphrates, by his host Sheik Mustapha, who
stated that the animal had been killed among the rocks near Sevi,
a small village about five hours down the river on the Mesopotamian
side ; it was the only specimen which he had ever seen. This Society
[the Zoological Society of London] has received more than one
specimen from Syria, and it is not improbable that the species may
be found in some parts of Asia Minor proper."
Tristram wrote in 1884 (p. 19) of this species in Palestine: "This
graceful Leopard is scarce, but still haunts the wooded hills of Galilee
and the neighbourhood of Tabor. East of Jordan it is far more com-
mon, and is much valued by the Arabs."
The Cheetah has now become very rare in Palestine. Yet it is
still pretty common in the southern steppe. Its use for the chase is
now quite outmoded. (Aharoni, 1930, p. 332.) It "still lives in the
Negeb, in Transjordania and rare specimens also persist in the Pales-
tinian mountains. The author saw many skins, sold by Beduins
from Beersheba." (Bodenheimer, 1935, p. 105.) More recently Pro-
fessor Bodenheimer writes (in litt., March, 1937) that the animal
is now on the verge of extinction or extinct and that nothing can be
done to preserve it in Palestine, but that perhaps there is still a
chance to do so in Transjordania.
In 1909 Carruthers (1935, pp. 60, 70) found Cheetah tracks on the
north side of the Jabal Tubaiq, Arabia, approximately 150 miles
east of the head of the Gulf of Akaba.
Turkestan Cheetah
ACINONYX JUBATUS RADDEi Hilzheimer
Acinonyx raddei Hilzheimer, Sitz.-ber. Ges. naturf. Freunde Berlin 1913, no. 5,
p. 291, 1913. (Based upon a specimen purchased in Merv, Russian
Turkestan (Turcoman S. S. R.).)
This Cheetah seems to occur in very small numbers in the southern
parts of Russian Turkestan. The animals of northern Persia and
northern Afghanistan may belong to the same form.
286 EXTINCT AND VANISHING MAMMALS
It is distinguished by its extremely thick, long fur; ground color
light brownish gray ; very large spots reaching to the toes ; tail long-
haired and very bushy, with five half -rings (one perhaps a complete
ring) at the end (Hilzheimer, 1913, p. 291). (Cf. Satunin, 1909,
pp. 254-256.)
"Only in the western portion of Turkestan have I met with this
species, and even there only on the low plains" (Severtzoff, 1876,
p. 49).
This is doubtless the commonest of the large cats in Transcaspia.
It is distributed through the whole region on the lowlands, along
the river courses, and on the mountains. Each year the Turkomans
bring young Cheetahs for sale into the cities and military posts.
Training the animals for the chase is unknown to them. (Radde and
Walter, 1889, p. 1012.)
The following data are from Ognev (1935, pp. 313-314) : The
Cheetah is found from time to time as far north as the Mangyshlak
Peninsula (Karelin, 1883). It inhabits the Kara Tau, the western
spurs of the Tian Shan, the lower Syr Darya, the Zarafshan Valley,
and the steppes between Zarafshan, Syr Darya, and Kizil Kum,
reaching an elevation of 600-1,000 feet (Severtzov, 1873). It also
occurs on the Amu Darya (Zarudny, 1915) and in Tajikistan. It is
particularly numerous along the Murgab, Tejend, and Sumbar
Rivers (Bilkewicz, 1918). The Caucasian Museum has specimens
from Merv, Kizil Arvat, and the Kopet Dagh. The Cheetah is also
recorded from Mazanderan, northern Persia (De Filippi) .
About 1884 two cubs were obtained in northeastern Persia near
the Turbat-shaikh-jami River, a tributary of the Hari Rud (Ait-
chison, 1889, pp. 56-57) .
The Cheetah is observed irregularly and in very small numbers
in Turkestan on the frontiers of Persia and Afghanistan. Some are
killed, but not every year. (W. G. Heptner, in litt., December, 1936.)
King Cheetah; Cooper's Cheetah
AdNONYX REX PoCOck
Acinonyx rex Pocock, Abstr. Proc. Zool. Soc. London, no. 283, p. 18, Mar. 1,
1927 ("Umoukwe [=Umvukwe] Range, N.W. of Salisbury, [Southern]
Rhodesia"); Proc. Zool. Soc. London 1927, pt. 1, p. 250, pi. 1, April 6,
1927.
FIGS.: Pocock, 1927a, pi. 1, and 19276, frontisp.; Dollman, 1929a, p. 3, fig.;
Maydon, 1932, pi. 103.
The range as well as the numbers of the King Cheetah are dis-
tinctly limited, and special protective measures seem to be neces-
sary in order to insure its survival.
It is similar in size and proportions to the Common Cheetah (A.
jubatus) but distinguished from it by a bold pattern of black
ORDER CARNIVORA: CARNIVORES 287
stripes and blotches, which are longitudinal on the dorsal area and
oblique or longitudinal on the flanks; legs blotched and spotted to
the feet; basal half of tail with two longitudinal stripes, distal half
with irregular transverse stripes; ground color mostly cream-buff;
belly white. Skin measurements: head and body, 4 feet 2 inches
to 4 feet 5 inches; tail, 2 feet 6 inches. (Pocock, 1927a, pp. 250-
252.) The King Cheetah is by some considered as a color mutation
of the Common Cheetah (G. M. Allen, MS.).
FIG. 32. King Cheetah (Acinonyx rex). After Pocock, 1927.
The species is known only from Southern Rhodesia. Its range
may lie wholly within the northern division of the Southeastern
Veldt District of Bowen (1933, pp. 256, 260) .
This superb new Cheetah was brought to scientific attention in
1926, by Major A. L. Cooper, who sent to the British Museum the
skin of an animal trapped by natives in the Umvukwe Range
(Pocock, 1927a, p. 245).
Cooper (in Maydon, 1932, pp. 335-336) gives the following ac-
count:
"That this animal was known for some time past is borne out
by the fact that, twenty years ago, mention used to be made round
camp fires by natives of a beast that was neither Lion, Leopard,
nor Cheetah, and ... I believe was referred to as the 'Mazoe
Leopard.' It was apparently commoner in those days than it is now."
The skin now in the Salisbury Museum was purchased from
natives, who stated that they had killed the animal in the Macheke
district. There were four or five in the troop.
H. M. G. Jackson reported a similar skin at the American Mission
288 EXTINCT AND VANISHING MAMMALS
at Utambara. "It was also an old native police sergeant of his,
who, when shown the skin, said he knew the animal, told us its
native name, and informed us of its habits, namely, that it is ex-
tremely shy, never attacked domestic animals except possibly a
young kid, and, when chased by dogs, never took to a tree as a
Cheetah occasionally does. . . .
"It was found that Mr. Watters, Native Commissioner at Bitika,
possessed two such skins. . . . These were presumably obtained in
his district. Apart from these two, I found Mr. Lacey of Salisbury
also had a specimen, . . . killed some twenty miles south of Salis-
bury. . . .
"This is the history of the discovery, if it can be described as
such."
Pocock reports (1927a, p. 246) that the animal whose skin was kept
at the Utambara Mission "was shot in the Melsetter District close
to the Portuguese Border. . . . The natives were not at all afraid
of it as they were of leopards, and would attack it armed only with
assegais." He says also (19276, p. 19) :
In the interests of the preservation of the new species of cheetah the
following probabilities cannot be too strongly insisted upon. All the avail-
able evidence suggests that the animal has a restricted range and is nowhere
plentiful. Its distributional area is within reach of Salisbury, an easily
accessible centre; and the publicity now given to the existence of so hand-
some an animal will surely be taken advantage of by sportsmen and traders.
All the big museums in the world will be eager for its skin, and every
zoological garden will want live specimens for exhibition. It will, therefore,
command a high price, whether alive or dead, and the result will be per-
secution by hunters and trappers on such a scale as to threaten its extinction
unless the authorities in Rhodesia at once take such steps as may be neces-
sary to protect it.
There seem to be no nature reserves within the known range of
the King Cheetah.
Barbary Lion. Le Lion de Barberie (Fr.). El Leon
berberisco (Span.)
LEO LEO LEO (Linnaeus)
Felis leo Linnaeus, Syst. Nat., ed. 10, vol. 1, p. 41, 1758. ("Africa"; type locality
subsequently restricted to Constantine, Algeria, by J. A. Allen, Bull. Am.
Mus. Nat. Hist., vol. 47, p. 222, 1924.)
FIGS.: Reichenbach, 1836, figs. 1-2; Geoffroy and Cuvier, Hist. Nat. Mammif.,
vol. 1, pis. 114-115, 1824.
The Barbary Lion survived well into the twentieth century but
is now extinct.
"Very large, dusky ochery, with the mane very thick and long,
extending to the middle of the back; and a thick and heavy mane
ORDER CARNIVORA: CARNIVORES 289
on the under-parts. In the female the inside of the fore-legs is
white." (Lydekker, 1908, p. 416.) The female is paler and smaller
than the male (Cabrera, 1932, p. 181).
This extinct race ranged from Tripoli through Tunisia, Algeria,
and Morocco. Apparently there are no definite records of Lions
within the present confines of Egypt during historic times. The
Lion of Senegal is described as a distinct race (Leo leo senegalensis
(J. N. von Meyer)), although a single race of Cheetah (Adnonyx
jubatus hecki Hilzheimer) is accredited to both Senegal and the
Barbary States.
Tripoli. "About two hundred years ago the lion was found quite
commonly in Tunisia. About the same time*, so far as records go, the
last lion was killed in the adjoining Pashalik of Tripoli, where the
animal now seems to be entirely extinct." (Johnston, in Bryden,
1899, p. 564.)
Tunisia. "Down to the time of the French invasion of Tunis,
in 1881, lions were still found in the extreme north-western part of
the Regency, close to the Algerian frontier. . . .
"What has brought about the extinction of this animal is less
the persistent attacks of French or Arab sportsmen than the opening
up of the forests and the settling down of the people since the French
occupation. The herds are now so carefully tended that the lion has
little or no chance of feeding on them, while the Barbary stag and
the gazelles have in that region become very scarce." (Johnston, in
Bryden, 1899, pp. 562-564.)
The last Lion of Tunisia was killed in 1891 at Babouch, between
Tabarka and Am-Draham. The species was common up to the time
of the French occupation. Doubtless owing to troop movements, it
then retired to the most remote massifs. It could not survive in
contact with civilization. (Lavauden, 1932, pp. 5-6.)
Two specimens in the Ley den Museum (one killed in Tunisia in
1823, the other in "North Africa" about the same period) are prob-
ably the only wild-killed Barbary Lions that are preserved in any
museum and can be studied at present (Cabrera, 1932, pp. 182-183).
Algeria. Pease (in Bryden, 1899, pp. 564-566) gives the following
account:
The North African lion was in bygone ages undoubtedly very numerous. . . .
The Algerian lion has become so rare that it may be said to be nearing
extinction. ... It lingers only in the country that might almost be
described as the Mediterranean littoral zone, though an occasional lion is
still shot or tracked in the interior, as far inland as the district of Soukarras,
and certain places in the Aures. [During 1892-95] I do not remember hearing
of more than three or four being obtained in the whole province of Con-
stantine. In the provinces of Algiers and Oran they may be said to be ex-
tinct. So long ago as 1862 General Marguerite wrote that ... in the province
of Algiers . . . the average number killed did not exceed three or four a
290 EXTINCT AND VANISHING MAMMALS
year. . . . General Marguerite relates, that during his eleven years the
Beni-Mahrez, a tribe not numbering more than 100 tents, lost on average
annually, 3 horses, 25 cattle, and 75 sheep from the depredations of lions
and panthers .... Before the French came, the Turks had encouraged the
Arabs to destroy them by freeing the two great lion-hunting tribes, the
Ouled Meloul and Ouled Cessi, from all taxes and paying liberally for their
skins. The French gave only 50 francs for a skin.
Between 1873 and 1883 the process of extinction is measured in Govern-
ment returns. The numbers killed for the whole of Algeria were, in the last
six years of this period, 1878, 28; 1879, 22; 1880, 16; 1881, 6; 1882, 4; 1883, 3;
(1884, 1) ; and for the decade-
Province of Algeria 29
" " Constantine 173
" " Oran
202
There are a few lions still left in the Province de Constantine, in the thick
forests between Soukarras and La Calle.
According to Johnston (in Bryden, 1899, p. 564), "Lions still
linger here and there in South-East and South-West Algeria."
On the other hand, Lavauden, an eminent authority on the North
African fauna, fixes (1932, p. 6) the date of the Lion's disappearance
in Algeria at about 1891, when the last one was killed in the region
of Souk-Ahras.
Morocco. To judge by the literature of several centuries ago,
Morocco was then a veritable country of Lions. At the middle of
the seventeenth century they still abounded on the Mediterranean
coast. Toward the end of the eighteenth century the range extended
to Cape Nun, Ifni. By the middle of the nineteenth century the Lion
had retreated from the entire Mediterranean littoral. According to
old native hunters, about 1880 not a Lion remained north of the
low Bu Regreg and Taza Pass. Some years later many forests of the
Middle Atlas served as a refuge for bandits, and this fact, together
with the civil wars of those times, contributed to the disappearance
of the Lions. Even in 1901 Lions were said to be frequent visitors
to the forest of Budaa, near Azru. According to the ex-Sultan Muley
Hafid, there remained in Morocco about 1911 only a few Lions,
which lived in the forests of the Zaian and the Beni Mguild. Ap-
parently they survived at least to 1922 in the Middle Atlas, and
it is probable that they inhabited the Grand Atlas likewise till a
comparatively recent date. They inhabited especially the wooded
mountains.
The Lion figures largely in the folklore of Morocco.
Its rapid disappearance from this country constitutes a very
curious problem. Unlike the Cape Province and Algiers, Morocco
remained wild and uncivilized up to a quarter of a century ago.
Its inhabitants are far from being a hunting people, and few Euro-
ORDER CARNIVORA: CARNIVORES 291
peans have ventured there owing to the lack of security. (Cabrera,
1932, pp. 186-190.)
Heim de Balsac (1936, p. 98) places the disappearance of the
species in Morocco in the decade 1900-1910 a considerably earlier
date than Cabrera's.
Utilization by the Romans. "There is ... little doubt that the
Romans drew their chief supply of lions for the arena and gladia-
torial combats from Mauretania and Numidia." Pliny speaks of
hundreds at a time being shown by Pompey and Caesar in the
Roman arena. (Pease, in Bryden, 1899, p. 564.) This bespeaks a
great abundance of Lions in North Africa at that period.
European Lion
LEO LEO subsp.
Of the Lion that still existed in Greece in classical times, no
remains seem to have been found. If suitable material were avail-
able, the modern systematist would probably find means of dis-
tinguishing it from any living Lion as well as from its Pleistocene
ancestor (Leo spelaeus) that once roamed over a large part of
Europe. Up to the present time, however, it apparently has not
received even a subspecific designation.
Meyer (1903, pp. 65-73) has provided a useful summary of our
knowledge of the European Lion. Herodotus (ca. 484-430 B. C.)
reports many Lions between the Achelous River in Acarnania and
the Nestus River in Abdera (Thrace) and states that during
Xerxes's march through Macedonia (480 B. C.) Lions killed some
of his baggage camels. Aristotle (384-322 B. C.) assigns the same
range to the Lion, but speaks of it as rare. By A. D. 80-100 it was
considered entirely exterminated in Europe, as a result of a gradual
retreat before man and his culture.
"The Greek name for the lion is very ancient, and this suggests
that it refers to an animal indigenous to the country. Although the
evidence is not decisive, it seems probable that lions did exist in
Greece at the time of Herodotus; and it is quite possible that the
representation of a lion-chase incised on a Mycenean dagger may
have been taken from life." (Flower and Lydekker, 1911, vol. 16,
p. 737.)
Evidently Elliot regarded this Lion as identical with the pre-
historic Cave Lion (Leo spelaeus). He writes (1883, text to pi. 1) :
"The Cave-Lion disappeared from Britain towards the close of the
Postglacial period, and is considered to have retreated gradually
from Europe and become extinct between 340 B. C. and A. D. 100.
The cause of this disappearance, according to Dawkins, was the
warfare carried on against it by the people of those periods, as
292 EXTINCT AND VANISHING MAMMALS
exhibited by the leonine remains found in the ancient dwelling
places of the Postglacial men in Aurignac and La Madeleine. This
is probably a correct supposition; for neither was the temperature
unsuited for its continued existence nor had the supply of food
failed."
Asiatic Lion; Indian Lion
LEO LEO PERSICUS (Meyer)
Felis leo persicus Meyer, Dissertatio inauguralis anatomico-medica de genere
Felium [Vienna], p. 6, 1826. (Persia.)
SYNONYMS: Felis leo goojratensis Smee (1833); Leo asiaticus Jardine (1834).
FIGS.: Trans. Zool. Soc. London, vol. 1, pi. 24, 1834; Jardine's Nat. Libr.,
vol. 15, Mammalia, pi. 11, 1842; Elliot, 1883, pi. 1, upper right-hand fig.;
Pocock, 1930, pis. 1-3.
This Lion, once widely distributed from Asia Minor, Palestine,
and Arabia to Persia and India, is now almost or entirely reduced
to a small remnant in the Province of Kathiawar, in western India.
There may be also a few solitary survivors in Persia and Iraq, but
this is doubtful.
"On the average, the Indian lion has a scantier mane than the
African and, curiously enough, ... a fuller coat, a longer tassel
of hair at the end of the tail, a more pronounced tuft of hair on
the elbow joints and a fuller fringe of hairs on the belly. In size,
there is little to choose between the two. . . . The largest recorded
measurement of an Indian lion is 9 ft. 7 in., of an African lion
10 ft. 7 in." (Anonymous, 1935, p. 123.)
Asia Minor. In 1878 or 1879 Sheik Mustapha informed Danford
"that five years ago a Lion appeared near Biledjik [on the Euphrates
toward the Syrian border], and after destroying many horses was
done to death" (Danford and Alston, 1880, p. 53). This is the only
definite locality record I have found for Asia Minor.
Syria, Palestine, and Arabia. "The Lion has long been extinct
in Palestine, and among the inhabitants there is no tradition of its
existence. Yet of its former abundance there can be no question.
It is mentioned about 130 times in Scripture .... Within the
historic period it was common in Syria, Asia Minor, and Greece.
... It seems to have disappeared altogether from Palestine about
the time of the Crusades, the last mention of it being by writers of
the twelfth century, when it still existed near Samaria. ... It can
scarcely be said now to exist in Asia west of the Euphrates, unless
in Arabia, the latest trace being that a few years ago the carcase of
one was brought into Damascus. . . . The Arabs state it is found
in Arabia." (Tristram, 1884, p. 17.)
According to the Old Testament, the Lion was found in Lebanon
ORDER CARNIVORA: CARNIVORES 293
and on the Jordan. The ancient writers (Xenophon, Aristotle,
Strabo, Pliny, etc.) speak of lion hunts in Syria and also in Arabia.
(Meyer, 1903, p. 71.)
"To-day, the nearest [to Palestine] wild habitats of the lion are
the jungles of the Upper Euphrates and several Arabian oases. But
even in those places it must be on the verge of extinction." (Boden-
heimer, 1935, p. 114.)
Iraq (Mesopotamia) and Persia. In the early part of the last
century Lions were noted fairly commonly along the Euphrates and
the Tigris. The explorer Layard hunted them with the Bakhtiyari
chiefs in Arabistan, whose sheep and oxen suffered from the Lions'
depredations. By the middle of the century Layard reported the
species as then found rarely on the Tigris as far north as Mosul,
but frequently below Bagdad. He adds: "On the Euphrates it has
been seen, I believe, almost as high as Bir .... On the [Jebel?]
Sinjar and on the banks of the Khabour [in the northeast of the
present Syria], they are frequently caught by Arabs. They abound
in Khuzistan [western Persia]." (Kinnear, 1920, pp. 33-35.)
By 1891, according to Sir Alfred Pease (Book of the Lion), the
"lion is no longer found in Asia Minor, but exists in Mesopotamia
and Arabistan, between Poelis, west of Aleppo, and Deyr [in the
present Syria] , and in the Euphrates valley ... ; it is also found in
the lower part of the Karun river but is nowhere plentiful." (Kin-
near, 1920, p. 36.)
Blanford writes (1876, p. 29) : "The lion at the present day is
found in Mesopotamia, on the west flanks of the Zagros mountains
east of the Tigris valley, and in the wooded ranges south and south-
east of Shiraz. It nowhere exists on the table land of Persia." To
this 0. St. John adds (in Blanford, 1876, pp. 30-31) : "Lions, which
are very numerous in the reedy swamps bordering the Tigris and
Euphrates, are found also in the plains of Susiana, the modern
Khuzistan, and extend into the mountain country south of Shiraz
as far east as longitude 53." Acorns of an oak (Quercus aegilopi-
folia) "feed the wild pigs whose- presence tempts the lion into the
mountains of Pars. . . . The little valley of Dashtiarjan, thirty-
five miles west of Shiraz, is notorious for the number of lions found
in its vicinity. . . . Dashtiarjan is ... a perfect paradise for
swine, ... so that the lions have plenty to eat .... Every year
some four or five adult lions are killed in Dashtiarjan or the neigh-
bourhood, and a few cubs are brought in to Shiraz for sale."
Edward Thompson (London Times, August 19, 1932?) gives the
following reports for Mesopotamia: a Lioness and cubs seen by an
Indian trooper near Ahwaz in 1917; a Lion cub brought through an
Arab village near Sanniyat in 1916; and one shot in the Wadi
294 EXTINCT AND VANISHING MAMMALS
marshes a year later. He also mentions a report received by the
Bombay Natural History Society of the continued existence of Lions
in the Pusht-i-Kuh range of western Persia.
"There are Persian Lions, and the last time a pair of them was
seen in the South of Persia by French and English Engineers (in
1928) .... The animals were carefully watched for several hours
and were seen by hundreds of people. ... I understand that the
Persian ruler takes keen interest in their preservation and they
are not allowed to be shot." (Hasan Abid Jafry, in Hit., August
17, 1933.)
The Persian Lion is a thing of the past. Firearms, whose use
increased during the World War, were more dangerous to the Lion
in Persia than in Africa. In 1923 the last of its kind was killed
south of Shiraz. Yet the people still express belief in the existence
of Lions. In the swamp and reed areas of the Euphrates and the
Tigris I have been able to find no more trace of the Lion. The opera-
tions here during the World War paved the way for its extinction.
Skins of Persian Lions are still found in some mosques. (Becker,
1934, pp. 439-440.)
The Lion may survive in the wilder mountains of Luristan and
Khuzistan in southwestern Persia (Bombay Natural History Society,
in litt., December, 1936) .
"The Syrians frequently used the lion motif as a frieze decora-
tion, and at Persepolis, thirty miles northeast of Shiraz, where the
magnificent ruins of the palace of Darius the Great may still be
seen, the lion as a decorative architectural motif was constantly used.
In the embrasures of some of the great doors of Persepolis the winged
lions were magnificently carved." (Vernay, 1930, p. 82.)
Afghanistan and Baluchistan. "There is no evidence to show
that the lion inhabited Afghanistan or Baluchistan within historic
times" (Kinnear, 1920, p. 37) .
"I was told, while in Duzbad, the frontier town on the Baluch-
Persian border, that the lion existed in Afghanistan seventy-five
years ago. This is mere heresay, but it sounds quite reasonable."
(Vernay, 1930, pp. 82-83.)
In 1935 Admiral Philip Dumas reported seeing a Lion at close
range near the Bolan Pass, south of Quetta in Baluchistan (Jour.
Bombay Nat. Hist. Soc., vol. 38, no. 2, pp. 381-382, 1935).
India. "Within the present [19th] century, distributed over
much of Central, West, and North-west India; but now confined
in that country to the peninsula of Guzrat, unless a last remnant
still maintains a lingering existence in the jungles bordering the
Sind River in Bundelkund, which I now consider doubtful" (Blyth,
1863, p. 182).
ORDER CARNIVORA: CARNIVORES 295
Kinnear (1920, pp. 37-39) writes:
[The Lion] was formerly found in Sind, Bahawalpur and the Punjab,
becoming extinct round Hariana, in the latter province, in 1842. It was
however extinct in Sind before that date and the last on record was shot
near Kot Deji in 1810. Exactly how far eastwards the lion was a regular
inhabitant we do not know, though there is a statement of one being killed
in the Palamaw district, Behar and Orissa, in 1814, but whether this was
merely a straggler or not, there is no evidence to show. The southernmost
limit appears to have been the Narbada. In 1832 one was killed at Baroda,
while further north it was comparatively common round Ahmedabad in
1836. Central India in these early days was one of the strongholds of the
lion and to give an idea of its numbers we may mention that Lydekker was
informed that during the Mutiny, Colonel George Acland Smith killed up-
wards of 300 Indian lions and out of this number 50 were accounted for in the
Delhi district !
The occurrence of the lion in Cutch is doubtfully recorded. The lion
probably was found in Cutch at one time but the records are not satisfactory.
Dates of extermination in other parts of India, according to Kin-
near, are: Damoh district (Saugor and Narbada territories), 1847-
48; Rewah (between Allahabad and Jubbulpore), 1866; Goona,
1873; Abu and Jodhpur (Rajputana), 1872; Deesa (Guzerat), 1878;
Palanpur (Guzerat), 1880.
A map, showing dates of extermination of the Lion in various
localities in India, is given by Pocock (1930, facing p. 661).
"A small number [of African Lions] were imported into Gwalior"
about 1890-1900, "but after a few years they became a pest, killing
not only the cattle of the natives, but also the natives themselves,
so that the African lions were all eventually shot out. Also, the
tigers of Gwalior are famous, and as tigers will not permit lions to
remain in their territory, they must have helped to kill off the lions."
(Vernay, 1930, pp. 81-82.)
"In India the lion is verging on extinction. There are probably a
very few still living in the wild tract known as the Gir in Kattywar,
and a few more in the wildest parts of Rajputana, especially South-
ern Jodhpur, in Oodeypur, and around Mount Abu." (Blanford,
1888, p. 57.)
"In 1893 ... a rough census was taken [in the Gir Forest],
and the number remaining was estimated at twenty-six, which sub-
sequent estimate raised to thirty-one. . . . There are now esti-
mated to be only twenty lions remaining in the Gir, of which eight
are cubs. (Lydekker, 1900, pp. 270-271, quoting from The Asian
newspaper of June 19, 1900.)
"It is only in the Province of Kathiawar, a small peninsula north-
west of Bombay, that the true Asiatic lion can still be found. Even
there it exists only in the Gir Forest, an area of four hundred square
miles in the State of Janagadh. . . .
296 EXTINCT AND VANISHING MAMMALS
"It is only a question of time before the lion will disappear even
from this district, although at present it is closely protected. The
number, which is roughly estimated at 200, is not increasing. The
inevitable diminution of the forest, in spite of the restrictions against
cutting, and the possibility of disease owing to the confined area,
mean ultimate extermination." (Vernay, 1930, p. 81.)
Economics and conservation. The Lions "commit considerable
havoc amongst the cattle, which are brought into the Gir for grazing
purposes during the greater part of the year, besides helping them-
selves liberally to the sambar, nilgai, spotted deer, and pig with
which the forest abounds. ... A large number of lions are kept
in captivity in the State gardens at Junagarh, where they breed
very freely." (L. L. Fenton, in Lydekker, 1900, pp. 410-412.)
"It is reasonable to suppose that the factors which exterminated
it in Europe, Asia Minor and Syria and have brought it to the verge
of extinction in Mesopotamia and Persia, even if they have not
already achieved that end, were the same as the factors which
exterminated it over almost the whole of the area it occupied in
India. In my opinion there is no reasonable doubt that the main, if
not the sole, factor in the case of Europe and southwestern Asia
was man. At all events it was most emphatically not the tiger. . . .
"It is not unlikely, in my opinion, that the Kathiawar stock is
deteriorating in size from inbreeding." (Pocock, 1930, pp. 641, 665.)
"I ... hear that the status of the Indian lion, as far as preser-
vation and numbers are concerned, is most satisfactory. They have
of late been overflowing from their original reservations in the
Gir Forests of Kathiawar and Junagadh State, and have made
themselves unpopular by cattle killing. Their numbers are esti-
mated to be not less than 150.
"It must be remembered that there is no wild life in their present
habitat on which they can prey, and they live almost entirely on
village cattle. There are, of course, far more cattle there, as in other
parts of India, than are economically desirable ; but ... if the local
native rulers were to withdraw their protection, the lion would
speedily disappear. There is, however, little danger of this happen-
ing, and the villagers at present cooperate loyally in the protection
of these animals; even to the extent, in a recent case, of pulling a
lion out of a well into which he had fallen, with no little risk to
themselves." (C. H. Stockley, in Hit., May 29, 1933.)
"Even in the Province of Kathiawar, where tigers do not exist
and where no struggle for supremacy between these two giants of the
tribe could have taken place, the lion was slowly driven from the
Barda and Aleche hills, from parts of Dhrangadra and Jasdan as a
result of human settlement and the progress of cultivation.
ORDER CARNIVORA: CARNIVORES 297
"The number of lions in the Gir is computed to be well below a
hundred." (Anonymous, 1935, p. 125.)
Cadell (1935, pp. 165-166) writes as follows:
The animals are easily enticed across the boundary [of Junagadh State]
by a succession of tie-ups. ... To our certain knowledge . . . twenty- two
animals have been so slain within the three seasons ending in 1934. . . .
If every year the State has the very real honour and pleasure of enter-
taining distinguished guests for a lion shoot, it is a distinction which costs
a good many thousands of rupees. There is also the steady annual cost of
the sums paid in compensation to villagers and herdsmen whose cattle have
been killed by lions. . . .
There were supposed to be less than a dozen [lions] in 1880 .... As a
result of the strict preservation during the [British] Administration [from
1911 to 1920] the number was believed to have increased to fifty .... It
has since been stated . . . that there are now two hundred lions. . . . My
own opinion ... is that there are not much more than 75 to 80. ...
The pressure on Junagadh of suggestions for invitations to shoot lions
is ... increasing year by year .... Unless an agreement is reached [to
limit the number shot in one year to some such figure as five or six], and is
faithfully observed, the danger of the disappearance of the lion from the
fauna of India, and consequently from its last home in Asia, is obvious.
Cape Lion. Leeuw (Boer)
LEO LEO MEJLANOCHAITUS Hamilton Smith
Leo melanochaitus Hamilton Smith, Jardine's Naturalist's Library, vol. 15,
Introd. to Mammalia, p. 177, 1846. ("Cape of Good Hope.")
FIGS.: Griffith's Anim. Kingdom, vol. 2, pi. facing p. 428, 1827; C. H. Smith,
1846, pi. 10; Harris, 1840, pi. 29; Pocock, 1931, p. 208, lower fig.
The Cape Lion was the first of the African subspecies of Leo leo
to become extinct. The last record for the Cape Province is ap-
parently 1858; for Natal, 1865.
"The species is of the largest size, with a bull dog head ; . . . large
pointed ears edged with black; a great mane of the same colour
extending beyond the shoulders; a fringe of black hair under the
belly; a very stout tail, and the structure in general proportions
lower than in other Lions" (C. H. Smith, 1846, p. 177).
Pocock (1931, p. 208) writes as follows concerning a mounted
specimen in the Junior United Service Club, London, which is "said
to have been killed near the Orange River about 1830, probably
. . . near Colesberg":
The mane is not only remarkable for its luxuriance, length and extension
over the shoulder, but also for its blackness. It is indeed wholly black except
for the tawny fringe round the face and a certain amount of the same pale
hue low down on the shoulder.
The elbow-tuft and tail-tuft are likewise big and black ; but the belly fringe,
long and thick behind, becomes gradually shorter and thinner and gradually
disappears in front of the chest.
The interest of this lion lies in its being, so far as I am aware, the only
298 EXTINCT AND VANISHING MAMMALS
representative, in this country at all events, of the now extinct race of
splendid liong which formerly inhabited Cape Colony. . . .
The former range and the date of the extermination of the handsome South
African race are alike unknown.
Pocock adds that Smith's type specimen appears to have been a
rickety captive.
Roberts (1929, p. 92) quotes from Paterson (1789) the following
measurements of a lioness from the southern part of Cape Province :
total length, 8 feet 9^ inches; tail, 3 feet; "height before," 3 feet
8 inches.
Owing to lack of material, the exact limits of the range of the
Cape Lion will never be known. For present purposes the Cape
Province and Natal will be considered to comprise the former range.
"Civilization's steady march in South Africa during the past
twenty years has considerably limited the range of the lion. The
vast herds of game upon which he depended for food being swept
away, he has been forced to retire into remoter regions. From much
of the South Africa of Gordon Gumming he has vanished com-
pletely and forever." (Kirby, in Bryden, 1899, p. 549.)
"With regard to past times Kolben (1731), states that lions
were not uncommon near Cape Town as late as 1707, Sparrman
(1785), Paterson (1790), Thunberg (1795), and Barrow (1801), all
met with these animals as soon as they got away from the imme-
diate neighbourhood of Cape Town especially on the karoo and in
Uitenhage. The last record I have met with of the occurrence of a
lion south of the Orange River is of one killed with assegais near
Commetjes Post on the eastern frontier in 1842, as noted by Hall.
General Bisset shot a lion in Natal in 1865, which is probably the
last record for that Colony." (W. L. Sclater, 1900, vol. 1, p. 31.)
"Their food . . . consists of the larger game, mainly antelopes
of all kinds, but also includes zebras, giraffes, and buffaloes. They
will kill the donkeys and cattle belonging to prospecting and hunt-
ing parties, and will raid Kaffir kraals when driven to it by hunger.
Man-eating lions are generally old animals with bad teeth." (Haag-
ner, 1920, pp. 69-70.)
"It is stated that a lion was shot on the Ingonyama Tributary
of the Tsomo River, Transkei, in 1858. One was reported from Port
Alfred in 1846, and one was killed by shot from a spring gun on the
farm Lombards Post near Southwell, near Bathurst, about 1850."
(Hewitt, as quoted by Shortridge, 1934, vol. 1, p. 80.)
The nearest areas to the former range of the Cape Lion, that are
still inhabited by some form of the species, are South-West Africa,
the Kalahari, and eastern Transvaal. The last-mentioned area
(especially the Kruger National Park) is the home of the Sabi
Lion (Leo leo krugeri Roberts) .
ORDER CARNIVORA: CARNIVORES 299
Manchurian Tiger; Siberian Tiger; Amur Tiger;
Mongolian Tiger
PANTHERA TIGRIS LONGIPILIS (Fitzinger)
Tigris longipilis Fitzinger, Sitz.-ber. Akad. Wiss. [Wien], math.-nat. Cl.,
vol. 58, pt. 1, p. 455, 1868. ("Korea and Japan through northern China,
Manchuria, Mongolia, and Dzungaria north to southern Siberia, and west
through northern Tatary, Bokhara, and northern Persia to Mount Ararat
in western Armenia"; type locality restricted by Lydekker (1901, p. 288)
to "Amurland." Cf. Harper (1940, p. 194).)
SYNONYM: Felis tigris var. amurensis Dode (1871).
FIGS.: Pocock, 1929, pi. 4, pi. F (upper fig.); Morden, 1930, p. 548, fig.;
Pocock, 1937, p. 770, fig.
This Tiger, whose principal range is in northern Manchuria and
southeastern Siberia, seems to be declining rather rapidly in numbers.
It is somewhat larger and has a longer and thicker coat than
the Bengal Tiger; ground color paler; stripes less pronounced and
tending to become brown on the flanks. It is said to reach a length
of 13-14 feet.
Siberia. The following information is from Ognev (1935, pp.
292-293). Radde (1862) found the species along the Argun River
near Ust Strielka and near Nerchinskiy Zavod. Baikov (1925)
places its northern limits at the Shilka and the lower Zeya and
Bureya Rivers. It is numerous in certain parts of the southern
Ussuri district. A specimen in the Zoological Museum of the Uni-
versity of Moscow was said to have been taken in 1828 near Ba-
lagansk, west of Lake Baikal (Severtzov, 1855). (This record can
not be definitely allocated as to subspecies.)
Ford Barclay (1915, pp. 225-228) gives the following account:
Careful inquiries made in the summer of 1899 along the present route of
the Siberian Railway, as far as Chita and Niertschinsk on the Amur and
thence east along that river as far as Khabarovsk, elicited practically no
information ....
At Khabarovsk . . . plenty of information was forthcoming, and many tales
were floating about of the depredations of these animals during the winter
in close proximity to, and even in one case within, the town itself. . . . The
best ground was reported to be in the neighbourhood of Irma, ... a little
more than half-way to Vladivostok, [where large numbers of Wild Pigs
attracted the Tigers.]
At Irma I learnt that a number of skins were undoubtedly brought in every
winter, but it was believed that in most cases their wearers had been accounted
for by poison. . . .
In 1899 it was still not uncommon to find fresh footprints of tiger on a
winter's morning in any of the outlying streets of Vladivostok ....
In the mountainous district between Harbin and Vladivostok a certain
number are poisoned by the natives every winter.
According to Sowerby (1923, p. 31) , this Tiger occurs throughout
the forested areas of the Amur and the Ussuri, into Primorsk in the
300 EXTINCT AND VANISHING MAMMALS
extreme east. It is said to be most plentiful in the Amur Province,
round the mouth of that river; it is also numerous in the Ussuri
Valley. Westward it probably extends almost to the Yablonoi
Mountains.
"North of Khabarovsk they are extremely rare in the East, though
I understand there are a few in the Bureya Mountains. We saw
the tracks of only one in the region of Troitskov. In the Ussuri
River region they seemed to be relatively plentiful. East of Bikin
the forest seemed to be well tracked with tiger trails, but one tiger
throughout the winter can make a lot of tracks. We secured three
tigers fifty miles east of Bikin, during the winter 1929-30. As far
as I could learn, these were all the tigers taken in this region during
that winter. That the tigers have been able to hold their own up
to now seems somewhat encouraging, though the present extended
lumber activities of the Soviet Government take many Russian
hunters into the forest. Previously about the only people who
hunted the tiger were the Tungus tribes, with their primitive traps
and snares. I, personally, am under the impression that it is only
a matter of time until the tigers are reduced to the point of ex-
termination.
"Tigers bring a big price in China, as medicine, but the hunter
has to cover a lot of territory and work hard to get even one animal."
(G. G. Goodwin, in litt., May 18, 1937.)
Referring to the Maritime Province, Sowerby says (1934c, p. 40) :
"Tigers of the long-haired species, whose skins are so valuable, were
being secured in greater numbers than before, for, whereas formerly
about ten of these great cats were killed in the province each year,
over twenty had been killed during the first three months of the
present year."
W. G. Heptner writes (in litt., December, 1936) that more than
ten are killed each year in eastern Siberia. Hunting is allowed
throughout the year. Protection is given, however, in the reserve of
Sikhota Alin.
"Schrenck (1859, pp. 95-96) reported Fells tigris from Sakhalin
as a rare winter visitor from continent, but his statement seems very
doubtful" (Kuroda,1928,p.226). Ford Barclay (1915, p. 225) could
find no evidence of its occurrence there.
Manchuria. In this country, says Sowerby, the Tiger is "the most
dreaded of the carnivores." He continues (1923, pp. 30-32) :
His thick winter coat fetches a high price in the fur-markets of the world,
being worth far more than those of the Bengal, Persian or Sumatran tigers.
Not only is his skin of value, but his whole carcass; for the Chinese believe
that the bones, blood, heart, and even the flesh of the tiger have medicinal
properties of rare power, and will pay a goodly price for decoctions brewed
by the apothecary that contain such ingredients as powdered tiger's knee-cap,
ORDER CARNIVORA: CARNIVORES 301
or clotted tiger's blood. The heart of the tiger is supposed to impart to the
consumer the courage and strength of the tiger itself.
On this account the tiger has been hunted till he is almost extinct in
most districts of North China, where once he was common, and now
survives, even in Manchuria where he was once plentiful, only in the more
remote and inaccessible forest areas, such as the Ch'ang-pai Shan, the Khingan
Mountains, or the more or less unexplored and thinly settled areas of the Amur
and Ussuri. . . .
Formerly the tiger was extremely plentiful in all the forested areas of
Manchuria. Indeed, it is said, they were so plentiful along the route of the
western portion of the Chinese Eastern Railway when under construction,
that they became a positive pest, killing and carrying off workmen, till a
regiment of Cossacks had to be sent to cope with the situation. . . .
In the forests of North Kirin and in Central and Western Heilungkiang
tigers are killed by the local hunters every winter.
The same author adds (p. 33) that the Russian hunters in Man-
churia track the Tiger down in the snow, camping on its trail and
following it for as much as ten days or a fortnight. The Chinese
usually employ traps, pitfalls, and poison.
Mongolia. In view of the fact that one of the names applied to
the present subspecies is "Mongolian Tiger," it is surprising to find
such a dearth of definite records from that wide country. According
to Ognev (1935, p. 292), Radde (1862) reported the Tiger from the
district of Uriankhai (the present Tannu-Tuva) , but later explorers
have not found it there. (The subspecies of this region has not been
determined.) Various other references in the literature to Tigers
in Outer Mongolia give no information as to specific localities.
Apparently the only likely areas for their occurrence are in north-
eastern and eastern Mongolia, along the Siberian and Manchurian
boundaries.
Korean Tiger; North China Tiger
PANTHERA TIGRIS COREENSIS (Brass)
Felis tigris coreensis Brass, Nutzbare Tiere Ostasiens, pp. 4-5, 1904. (Korea.)
(Fide Kuroda, 1938, p. 40.)
FIGS.: Ford Barclay, 1915, pis. 84, 85; Sowerby, 1923, pi. 2 (coreensis^);
Sowerby, 1933, pi. facing p. 166; Ognev, 1935, pp. 285-286, figs. 129-131.
This Tiger apparently occurs in small numbers from Korea and
southern Manchuria westward through the eastern border of Inner
Mongolia and through North China. Its southern limits, where it
presumably intergrades with the South China form, are not definitely
known but perhaps may be roughly fixed at the divide between the
Hwang Ho and the Yangtze Kiang Basins.
The North China form differs from the Manchurian Tiger "in
being smaller,- much darker and more fully striped and in having a
shorter less woolly winter coat" (Pocock, 1929, p. 531).
302 EXTINCT AND 'VANISHING MAMMALS
Korea. "In the Korea great value is apparently placed upon the
skins, which are reserved for the chiefs" (Elliot, 1883, text to pi. 3).
Ford Barclay (1915, pp. 228-231) gives the following account:
Tiger are probably more numerous in the north than in the southern part
of Korea ....
In the neighbourhood of the foreign mining concessions, near the Yalu,
dynamite is or was used with some success by native hunters, a small, specially
constructed bomb being somehow concealed in the bait. Lately, however.
. . . the Japanese police have forbidden the supply of dynamite for this
purpose. Drop traps, weighted with stones and huge logs, are very common,
and many tigers are accounted for in this way every year.
[In Manchuria] the natives lay down poison wholesale. This is forbidden
now in Korea ....
My own most successful hunts have been in the island of Chindo, . . .
situated at the south-west corner of Korea. . . . Early this year (1914) the
body of a tiger was washed up on the west coast of Japan south of Matsue,
at least 120 miles from the nearest mainland, from whence alone it could have
come; yet, as reported in the press, its condition was such that the skin was
removed for dressing and parts of the flesh sold for consumption 1 . . .
This demand for tiger flesh on the part of the Japanese is a curious survival
of barbaric superstition in such a highly civilized race. One of their chief
officials sent me an urgent request for a shoulder on hearing of a successful
hunt. This joint for some reason is supposed to possess greater medicinal
virtue than any other, and the shoulder blade ground to powder is a certain
cure in the most advanced stages of insanity!
When a tiger is killed [in Korea] notice is at once sent to the elders of all
villages within a radius of five miles, [and on their arrival a] wrangle ensues
as to who are to be the privileged half-dozen to partake of a cupful of the
ambrosial liquid left in the abdominal cavity, after the removal of the
intestines. . . .
Among both Chinese and Koreans, tiger's blood is believed to have an
extraordinarily rejuvenating effect, greater even than the highly prized wapiti
or sika horn ....
Of the twenty odd skins I have seen in South Korea all have been much
darker in colour than the half-dozen brought for my inspection in East
Siberia ....
In the happy days before the Japanese occupation and the consequent
confiscation of fire-arms, when the depredations of a tiger became too pro-
nounced, the active male inhabitants of the villages in the neighbourhood,
perhaps half a dozen, armed with matchlocks, and as many more with heavy
spears, would arrange for a day or two's driving in the adjacent hills. Occa-
sionally these hunts were successful.
"In North Corea tigers are said to be still fairly numerous, and
every year some are killed there by sportsmen" (Sowerby, 1923,
p. 31).
In 1922 Kermit Roosevelt (in Roosevelt, Roosevelt, Derby, and
Roosevelt, 1927, pp. 41-84) undertook an extensive but unsuccessful
Tiger hunt with beaters in various localities of northern Korea. Some
old tracks were found, but apparently the species is by no means
common there.
Manchuria. Sowerby 's records (1923, p. 30, pi. 2) from the
ORDER CARNIVORA: CARNIVORES 303
Ch'ang-pai Shan, close to the Korean border, may refer to the
present subspecies rather than to longipilis.
Inner Mongolia. "A stuffed tiger's skin used to repose in a temple
in Lama Miao (Dolonor) .... It was said that the animal . . .
was killed in the streets of Lama Miao itself, having wandered from
the Wei-ch'ang, or Hunting Grounds, to the east of that town."
(Sowerby, 1923, p. 32.)
China. Sowerby (1923, pp. 31-32) writes of this Tiger in China:
How far west it extends is difficult to say, but it certainly reaches the
western border of the province of Shansi, in North China, and southward
reaches at least to the middle of the southern half of that province. From
there it extends northward into Mongolia and in a north-easterly direction
through Chihli, where it still occurs in the wilder parts of the Tung Ling
and Wei Ch'ang (the Eastern Tombs, and Imperial Hunting Grounds) to
the North-east and North of Peking ....
In North China the tiger is becoming increasingly rare. In 1909 I saw the
tracks in the snow of what must have been a very large animal in the
mountains of West Shansi, in the Ning-wu district. I also heard of tigers
in the Ko-lan Chou area and the Chao-ch'eng Shan, both heavily forested
districts further south in the same province. Further south still near P'ing-yang
Fu a tiger was killed by the natives about the year 1912. I have seen skins
of tigers that were said to have come from the Kuei-hua Ch'eng area in North
Shansi, and they were undoubtedly of the true long-haired type. The natives
in this area also insisted that tigers occurred there. . . .
According to Chinese accounts tigers also exist in Kansu, and on the
Thibetan border, but I have been unable to get any satisfactory verification
of this. It is more than likely that these animals occur for a considerable
distance west of Kuei-hua Ch'eng into that little known mountainous country
leading to the Ali Shan.
In the early part of the present century an old native hunter
reported the occurrence of three Tigers in the Eastern Tombs forest,
in Hopei, during his lifetime. In 1932 a Tiger was killed after it
had invaded a shop in the Yu Hsiang district of South Shansi.
(Sowerby, 1933, pp. 167-168.)
Owing to lack of specimens, it has not been determined whether
the occasional Tigers reported in Szechwan (cf. Wilson, 1913,
pp. 178-179, and Weigold, 1924, p. 74) belong to the North China
or to the South China form. ^
"Tiger-bones ... are a highly prized Chinese medicine, and are
supposed to transmit vitality, strength, and valour to those who
partake of them. In the Imperial Maritime Customs Trade Returns
of Hankow for 1910 is the following item: 'Tiger-bones, 77 piculs;
value, Tls. 6522.' " (Wilson, 1913, p. 179.)
"It is problematical whether or not predatory animals should
be protected in a thickly populated country like China, but it seems
a pity that such fine carnivores as the Chinese tiger (Panthera tigris
styani Pocock), the Amoy tiger (Panthera tigris amoyensis Hilz-
304 EXTINCT AND VANISHING MAMMALS
heimer) [,] the Manchurian tiger (Panther a tigris amurensis, Dode)
. . . should not be preserved as part of this country's wonderfully
rich mammalian fauna" (Sowerby, 1937, p. 257).
[In South China, from Chekiang and Hupeh southward, and also
westward to Yunnan, the Tiger occurs somewhat more commonly
than in the more northerly regions. To this form Hilzheimer has
given the name of Felis tigris var. amoyensis (Zool. Anz., vol. 28,
p. 598, 1905; type locality, presumably the vicinity of Hankow,
Hupeh) . It is recognized by G. M. Allen (1938, p. 480) , who regards
Panthera tigris styani Pocock (1929) as a synonym. W. L. Smith
(1920, pp. 355-363) gives an extremely interesting account of the
methods of the native hunters in the vicinity of Amoy, who, armed
only with torches and trident spears, track the Tigers into caves.
There is also an account of the Tiger of Fukien by Andrews (in
Andrews and Andrews, 1919, pp. 44-66) .
In French Indo-China, Siam, and the Malay Peninsula the Tiger
seems to be moderately common. For example, the number in Cochin
China is estimated at 200-300; here it is of interest from the point
of view of big-game hunting, but not commercially (Roche, Chef
du Service Veterinaire du Cochinchine, in litt., 1937) . Rodolphe M.
de Schauensee informs me that the Tiger is common in Siam (Thai-
land) but preys chiefly on the wild game and does not seem to be
regarded as a serious pest. The Tiger of these regions is not dis-
tinguished by Pocock (1929, pp. 532-533) from the Indian Tiger.
The Indian or Bengal Tiger (Panthera tigris tigris (Linnaeus))
ranges westward through Burma to India, where it inhabits the
greater part of the Peninsula from the southern slopes of the
Himalayas southward, but avoids the treeless and desert areas. It
is not yet rare enough to call for any special discussion in this
report.]
Tiger of Chinese Turkestan
PANTHERA TIGRIS LECOQI (Schwarz)
Felis tigris lecoqi Schwarz, Zool. Anz., vol. 47, no. 12, p. 351, 1916. ("Gebiet
von Kurla, Lop-nor-Gebiet" (probably near Bagrash Kul), Chinese
Turkestan.)
All the Tigers of Chinese Turkestan will be treated for convenience
under this name, although the exact limits of the subspecies are
unknown. Evidently the animal is not at all numerous, and its
numbers may have declined to the point of extinction. In his review
of the Tigers, Pocock (1929) seems to have overlooked the name
of this subspecies as well as the occurrence of any Tiger in Chinese
Turkestan.
ORDER CARNIVORA: CARNIVORES 305
This is a very brightly colored Tiger, with regular pattern, fore-
legs unstriped in front, conspicuous shoulder tufts, short neck mane,
thick cheek whiskers, slightly lengthened abdominal hair, and a thick
winter pelage. It differs from the form of Russian Turkestan in
having smaller stripes and dull brown instead of black thigh mark-
ings. (Schwarz, 1916, p. 352.)
J. H. Miller (in Carruthers, 1913, pp. 582, 609-610) writes of this
Tiger:
The dense jungles which cover so large a portion of the [Dzungarian]
lowlands . . . are the haunts of the tiger ....
The tiger inhabits the same country as the wapiti, though, perhaps, keeping
rather more to the dense reed-jungle. It is, however, not entirely restricted
to the plains, for in the Kash, Kunguz, and Jingalong valleys, on the Upper
Hi River, it is found at an altitude of from 4,000 to 5,000 ft. among the
thick scrub on the edge of the spruce forest. Every year a few tiger-skins
find their way into the Urumchi, Manas, or Shi-Kho bazaars. They are, in
nearly every case, secured in winter, by the farmers and herdsmen living
on the edge of the jungle, by means of poisoned carcasses of sheep or goats.
Very few of the natives would dare to fire at a tiger .... Wild-pig . . . are
undoubtedly the tigers' staple food, but during the winter they occasionally
raid a farmer's flocks, and it is then that poisoned carcasses are laid out for
them. . . .
I doubt if they are anywhere numerous. . . .
It must be remembered that the tiger which inhabits Dzungaria and the
Tarim basin, also the Ala Kul, Balkash, Syr Darya, and other portions of
Russian Turkestan, is a very different animal to the Manchurian variety.
It is not so long-haired, and it is considerably smaller and less finely marked.
Theodore Roosevelt (in Roosevelt and Roosevelt, 1926, p. 166)
writes of Tigers in the Tian Shan: "We were told that they existed
no longer in the Tekkes. . . . They [the natives] said that during
the last ten or fifteen years the native hunters had killed them off
with poisoned meat."
"The tiger . . . formerly ranged in the forests on the edges of the
Tarim Basin and the swampy areas along the northern slopes of the
Thian Shan. . . . The tiger seems to have been exterminated.''
(Morden, 1927, p. 123.)
Alpheraky (1891) reported the species from the Tekes and the
lower Kunges, tributaries of the Hi River in Dzungaria (Ognev,
1935, p. 291).
"The . . . tiger, which formerly inhabited the woods of the middle
Tarim, seems to be dying out" (Hedin, 1940, p. 149) .
Caspian Tiger; Persian Tiger
PANTHBRA TIGRIS VIRGATA (Illiger)
Felis virgata Illiger, Abhandl. K. Akad. Wissen. Berlin, 1804-11, physikal.
KL, pp. 90 and 98, 1815. ("In Persien und am Kaspischen Meere"; type
locality restricted by Harper (1940, p. 194) to the "Province of Mazanderan,
northern Persia.")
11
306 EXTINCT AND VANISHING MAMMALS
SYNONYMS: Felis (Tigris} tigris septentrionalis Satunin (1904); Felix tigris
trabata Schwarz, 1916.
FIGS.: Heck, Lebende Bilder, p. 157, 1899; Kennion, 1911, pi. facing p. 251;
Pocock, 1929, pi. D, lower fig., pi. 3; Ognev, 1935, figs. 121-124.
While the Indian or Bengal Tiger (Panthera tigris tigris x ) prob-
ably exacts a greater annual toll in human lives than any other
carnivorous mammal, some of its races in western, central, and
eastern Asia and in the Malay Archipelago conflict to a far less
degree with the interests of mankind. In any event, their numbers
have been reduced to a point where they are entitled to a place
in the present work.
The Caspian race is "generally a medium sized or smallish tiger
with a thick longish winter coat, dark in colour, with numerous,
close-set stripes showing a marked tendency to brownness on the
whole or parts of the body." Length of male, about 10 feet 8 inches;
of female, about 8 feet 6 inches. (Pocock, 1929, pp. 522, 540.)
The range of this Tiger extends from Transcaucasia (formerly)
through northern Persia to northern Afghanistan; presumably the
same form occurs northward to the Aral Sea and Lake Balkash in
Russian Turkestan (formerly to the upper Ob Basin and the Altai
region) .
Transcaucasia. "A few are annually killed in Turkish Georgia"
(Blyth, 1863, p. 182).
Satunin reports (1906, pp. 308-309) as follows on the Tigers of
Talish:
At the time of Radde's first expedition to Lenkoran in 1866 Tigers
were still very numerous there. In seven weeks six fresh skins were
offered him. But in 1879-80, in the course of eight months, he
could not secure a single fresh skin. Tigers still occurred, but were
very rare. According to the hunters' reports, the animals were quite
extirpated somewhat later, but in the 90's they began to increase,
and at the time of my expedition (1897-99) two to four specimens
were taken annually. At present they occur chiefly in the Prisib
district of Lenkoran, both in the lowland forests and in the foot-
hills. In 1899 Tiger tracks were found on the Mugan Steppe, where
the animal had gone apparently in pursuit of Wild Boars.
Satunin also expresses here the conviction that the numerous
reports of Tigers in other localities of Transcaucasia are due to
a confusion of this species with the Leopard. In a previous paper
(1896, pp. 289-290) he had stated that they occurred formerly as
far as the ridge of the Great Caucasus, and he had quoted Nord-
mann's report of Tigers killed near Tiflis in 1835. The species is
now exterminated in Transcaucasia (W. G. Heptner, in litt., Decem-
ber, 1936).
i Fells tigris Linnaeus, Syst. Nat., ed. 10, vol. 1, p. 41, 1758. (Bengal.)
ORDER CARNIVORA: CARNIVORES 307
Persia. Gmelin (1774, vol. 3, pp. 485-486) reports the animal
as pretty common in the forested mountains of Mazanderan. It
seldom makes unprovoked attacks on man. The skin is highly
prized, and is used for a horse-covering.
Blanford writes (1876, p. 34): "The tiger is only found in
Persia in the Caspian provinces, Mazandaran, and Ghilan, lying to
the north of the Elburz mountains .... These provinces, unlike
the plateau of Persia, are covered with dense forest, and in them
the tiger ranges up to an elevation of at least 5000 or 6000 feet."
To this St. John adds (in Blanford, 1876, p. 34) : "Tigers are
very numerous in the Caspian provinces of Persia, and in the
Caucasus as far as the mouth of the Araxes. . . . Cubs are often
captured in Mazandaran and brought to Tehran. I have seen speci-
mens in the Bagh-i-Washi quite equal in size to Bengal tigers."
In Mazanderan, sometime prior to 1911, "Col. Kennion only came
across two examples of this tiger; and there is reason to fear that
the race is on the wane" (Pocock, 1929, p. 522). "Considering the
abundance of game and the fewness of the tigers' foes, it is quite a
problem why the latter are not more numerous in these parts"
(Kennion, 1911, p. 246).
The British Museum has a specimen obtained at Astrabad in 1882
or earlier. In Astrabad and the adjacent portion of Turkestan the
Tiger occurs in various localities, including the Gurgan, Atrek, Sum-
bar, and Chandir Rivers (Ognev, 1935, pp. 289-290).
Afghanistan. In this country, as in Persia, the species appears
to be restricted to the northern part. "The tigers of the Perso-
Turkestan district . . . were doubtless excluded from India by the
Hindu Koosh and the desert areas of Persia and Baluchistan"
(Pocock, 1929, p. 509).
"Ferrier in his 'Caravan journeys' speaks of tigers in the jungles
of the Hari Rud north-west of Herat" (St. John, in Blanford, 1876,
p. 34).
The Afghan Delimitation Commission (1884-85) obtained a speci-
men from Karaol-khana on the Murgab close to the Turkestan
boundary. Tracks were reported in the valley of the Hari Rud,
and were also found at the Chashma-sabz Pass, at an elevation
of 5,000 feet, in the Paropamisus Range. "During summer . . .
they wander over the great rolling plains of the Badghis [on the
north side of the Paropamisus Range], ascending to higher altitudes
with the increase of heat, depending for their food on Pig, Oorial,
and even Ibex. In winter they resort to the . . . thickets of the
larger streams and main rivers, to which their usual food, the Pig,
also retires. The Turkomans say that an old and toothless Tiger is
especially destructive to sheep." (Aitchison, 1889, p. 56.)
308 EXTINCT AND VANISHING MAMMALS
Russian Turkestan and Western Siberia. Ehrenberg reports
(1831, p. 389) that Tigers are frequently observed on the Tarbagatai
Mountains southwest of Zaisan Nor; also that the Cossacks of the
Irtish have several times killed Tigers on the Kirghiz Steppe, spear-
ing them from horseback.
"North of the Hindu Kosh, Tigers occur in Bokhara, and proved
troublesome to the Russian Surveying Expedition on the shores of
the Aral in midwinter. They are also found on the banks of the
Irtisch, and in the Altai region." (Blyth, 1863, p. 182.)
Atkinson (1858, p. 282) mentions four specimens in the museum
at Barnaul, western Siberia. "The tigers were killed in Siberia at
different places, some at a distance of about five hundred versts
from Barnaoul ; they had come from the Kirghis Steppe, and crossed
the Irtisch into the Altai in the region around Bouchtarminsk. . . .
They are rarely found in Siberia; it is only when they are driven
from the steppe by hunger that they cross the Irtisch most prob-
ably when following the track of their prey : many peasants do not
even know them by name." Atkinson also reports (p. 486) many
Tigers about the western end of the Ala Tau, southeast of Lake
Balkash.
According to Severtzoff (1876, p. 49), the Tiger "is common in
Turkestan, especially up to about 4000 feet altitude; but beyond
that it is rare in winter, and only in the summer does it visit localities
which are higher than 7000 feet."
Carruthers writes (1915, pp. 149-150) : "In the same locality
[Oxus or Amu Darya Valley] inhabited by the Bokharan stags,
tigers are fairly numerous. These we know range the whole course
of the Oxus from the Sea of Aral to the foot of the mountains near
Kulab. They are seldom hunted or seen. I have good reason to
believe they wander across the desert from the Oxus to the lower
Zarafschan. The natives speak of them, and I am certain I heard
one one night in the saxaul forests which surround the swamps
where the river loses itself in the sands, and where large numbers of
wild pig roam."
The British Museum has a skull from the vicinity of Find j eh,
on the Murghab (Pocock, 1929, p. 522).
In Turkestan the Tiger reaches its northwestern limit at the
Gulf of Karabugas on the Caspian Sea, avoiding the Ust Urt Plateau.
It was formerly numerous on the Murghab and Tejend Rivers, the
last having been killed in that region in 1904. During a period of
some years prior to 1915 nine Tigers were killed in the Syr Darya
region. The species also occurs in the valley of the Chu and on the
Amu Darya delta. In 1887 it was reported as abundant on the lower
Hi River and on the southeastern shore of Lake Balkash; by 1930
its numbers in this region were few. There are old records from the
ORDER CARNIVORA: CARNIVORES 309
Tarbagatai Mountains southeast of Zaisan Nor, and from Zmeino-
gorsk, Bisk, and Barnaul in the Ob Basin of western Siberia. The
Tiger has entirely disappeared from its former haunts in the Dzun-
garian Alatau. In Tajikistan it occurs on the upper Vashni and on
the Kafiringan Darya. (Ognev, 1935, pp. 273, 290-292; map, p. 295.)
The following information is from W. G. Heptner (in litt., Decem-
ber, 1936) : The Tiger is found in limited numbers, but regularly,
at the mouths of the Amu Darya and the Syr Darya and on Lake
Balkash. It is more common on the upper course of the Amu Darya
and on its right tributaries. It comes over accidentally from Persia
and Afghanistan to Kopet-Dag, the upper Tejend, the upper Mur-
ghab, and Transcaucasia (Talish) . During the past 50-70 years the
numbers have been considerably reduced by hunting. The range
has also been reduced, and in certain areas (Transcaucasia, middle
Syr Darya, and Murghab) the Tiger is now exterminated. It is
difficult to estimate the total number, but there may not be more
than 200 in Russian Turkestan. The best areas are the headwaters
and the mouth of the Amu Darya. The Tiger is rarely met with
at the mouth of the Hi River on Lake Balkash, where probably only
ten or twelve animals exist. At the mouth of the Syr Darya it is
probably only a visitor, coming from Amu Darya. Hunting is
allowed throughout the year.
Javaii Tiger ,
PANTHERA TIGRIS SONDAICA (Fitzinger)
Tigris sondaica Fitzinger, 1 Sitz.-ber. Akad. Wiss. [Wien], math.-nat. Cl., vol. 68,
pt. 1, p. 454, 1868. ("Java und Sumatra"; type locality restricted by
Schwarz (1912, p. 324) to Java.)
The meager information available concerning the status of the
Javan Tiger indicates that it is no longer very numerous or generally
distributed on that island.
"Ground-colour light rusty; stripes very narrow, often duplicated.
. . . Fur short and close." (Schwarz, 1912, p. 325.) "Apparently
closely resembling the Sumatran race in size and coloration, but
distinguished from it, and from all other tigers, by the marked con-
striction of the occiput" (Pocock, 1929, p. 541).
In 1851 Horsfield (p. 44) remarked that Tigers were "numerous
and destructive ... in many parts of Java."
"Many tigers . . . may be found" on the Oedjoeng koelon Penin-
i This name is antedated by Felis tigris sondaicus Temminck (Coup-d'oeil
Possessions Neerlandaises, vol. 2, p. 88, 1847). It is highly questionable, how-
ever, whether Temminck's excessively brief and insufficient description ("le
grand tigre raye de Sumatra et de Java forme une espece distincte du tigre raye
du continent de FInde") is nomenclaturally valid.
310 EXTINCT AND VANISHING MAMMALS
sula, at the extreme western end of Java, which constitutes a nature
reserve (Dammerman, 1929, p. 34) .
"In Java, the tigers living up to 1914 in the swamp country near
Maoek, are now extirpated. In 1931 they were seen on the Goenoeng
Malabar. They are also found in the Baloeran District, southern
Banjoewangi, in the Southern Mountains, and near Banjoemas. Two
to four are shot every year at Tampomas. Finally a number of
tigers are also reported from S. E. Garoet" and from the Midangan
district. (Heynsius-Viruly and Van Heurn, 1936, p. 58.)
[The Sumatran Tiger (Panthera tigris sumatrae *) , although less
common than formerly, is still numerous in various districts, and its
protection is not urged at present. (Heynsius-Viruly and Van Heurn,
1936, p. 59).]
Bali Tiger
PANTHERA TIGRIS BALICA (Schwarz)
Felis tigris balica Schwarz, Ann. Mag. Nat. Hist,, ser. 8, vol. 10, p. 325, 1912.
("Den Pasar, Siid-Bali.")
FIG.: Schwarz, 1913, p. 71, fig.
In Bali the Tiger seems to be in rather imminent danger of
extinction.
It is very similar to the Javan Tiger, but smaller; ground color
somewhat brighter, and the light markings clearer white; fur short
and close. Head and body, 1,530 mm.; tail, 580 mm. (Schwarz,
1912, p. 326.)
About 1909-12 the Tiger was considered fairly common in Bali;
yet information concerning damage done by it was not forthcoming
(Schwarz, 1913, p. 73).
"A few yet live in West Bali, but they are having a hard time
because they are much sought by hunters from Java, so that they
will certainly disappear within a few years. The species also exists
in N. W. and S. W. Bali." (Heynsius-Viruly and Van Heurn, 1936,
p. 58.)
Order PROBOSCIDEA: Proboscideans
Family ELEPH ANTID AE : Elephants
The Elephants are composed of an Asiatic genus (Elephas) and
an African genus (Loxodonta) . Lydekker (1916) recognizes 4
Asiatic forms and 11 African forms, but Dr. Allen (1939b) ques-
i Panthera tigris sumatrae Pocock, Jour. Bombay Nat. Hist. Soc., vol. 33.
no. 3, p. 535, pi. H, 1929. ("Deli, Sumatra.")
ORDER PROBOSCIDEA: PROBOSCIDEANS 311
tions the taxonomic status of all but 4 in the latter group. The
distribution of the family covers southeastern Asia (India, Burma,
Ceylon, Thailand, French Indo-China, Malay Peninsula) , Sumatra,
Borneo (introduced?), and the greater part of Africa south of the
Sahara. Accounts of three forms are supplied herein.
Malay Elephant
ELEPHAS MAXIMUS HIRSUTUS Lydekker
Elephas maximus hirsutus Lydekker, Abstr. Proc. Zool. Soc. London, no. 130,
p. 20, 1914. ("Kuala Pila district of the Negri Sembilan province of the
Malay Peninsula" (Lydekker, 19146, pp. 285-286).)
FIGS.: Lydekker, 19146, p. 285, fig. 1; Lydekker, 1916, vol. 5, p. 84, fig. 25.
The Elephant of the Malay Peninsula is regarded by competent
authorities (e. g., F. N. Chasen, in litt., March 31, 1937) as "a
vanishing form."
This subspecies is "characterized by the square, instead of tri-
angular, form of the ear, the early date at which its upper margin
is bent over, and the presence in the young condition ... of a
thick coat of black and in part bristly hair" (Lydekker, 1914a,
p. 20).
The northward range of the Malay Elephant has not been de-
termined; it will here be provisionally considered to extend as far
as the Isthmus of Kra, in Peninsular Siam. In the remainder of
Siam and in French Indo-China the Elephant belongs presumably
to the Indian subspecies and is reported as more or less common
(Gyldenstolpe, 1919, p. 169; James L. Clark, in litt., June 26, 1936;
P. Vitry, in litt., December, 1936; Roche, in litt., 1937).
Malay States Flower says (1900, p. 365) : "Wild elephants do
not occur in either Penang or Singapore, nor are tame ones em-
ployed there; but on the continent, both in Siam and the Malay
Peninsula, elephants are found wild in suitable localities, and are
trained for various purposes. ... I saw more or less trained
elephants in ... Kedah, and Perak, but in the Southern Malay
States the people do not seem to catch and tame them." He also
(p. 366) quotes H. J. Kelsall (1894) to the effect that "the elephant
appears to be common throughout Johore"; and H. N. Ridley
(1894) as remarking that "the elephant, though common all through
Pahang, is never caught and tamed."
Referring to conditions from 1900 on, Burgess writes (1935,
p. 249) : "Elephants roam all over the peninsula and are common as
far south as Johore. . . . Since only a small fraction of the jungle
has yet been cleared, the probabilities are that large herds have
not been seen."
312 EXTINCT AND VANISHING MAMMALS
Hubback (1923, pp. 24-25) reports on "the damage done to plan-
tations by elephants" in the Malay Peninsula:
It is a very extraordinary thing, but all wild animals which browse seem
to acquire an unholy craving for the bark and leaves of Hevea brasiliensis.
Wild elephants especially, once they have tasted the bark, seem to go mad
for it. I have absolutely trustworthy evidence of an eye-witness who has
seen elephants strip the bark, from rubber-trees by first catching hold of a
small piece with the tip of the trunk and then pulling upwards, so that
a strip of bark is taken off the tree. In an incredibly short time the tree is
ruined. Then they love to lean against the trees, and I suppose are sur-
prised and annoyed when they fall over. Undoubtedly these wild elephants
are in certain places a serious nuisance.
A further account is given by Hubback in the Report of the Wild
Life Commission of Malaya (vol. 2, 1932) . He says that elephants
are not uncommon south of Gunong Sinting, between that mountain
and the Pahang border, and continues:
It is a fact beyond question that wild elephants do and have done consider-
able damage amounting to values of thousands of pounds. Had it not been
for elephants in Malaya still larger areas planted with rubber would now be
yielding latex. These facts are not in dispute. . . .
The elephants known as the "Carey Island Herd," which lived on a large
island on the coast of Selangor, which island was given out for agriculture,
were all ultimately destroyed. Their death warrant was really signed when
the grant for the land was made out. Then there is the "Kuala Selangor
Herd" which has been almost totally exterminated; a cow and a calf being
reported as the sole survivors. The destruction of the survivors was advo-
cated. This herd must have consisted of 40 or 50 animals thirty years ago.
The "Labu Herd" in Negri Sembilan has been practically eliminated. In
Lower Perak the "Chikus Herd" of elephants has given a lot of trouble and
many of them have been shot. In many other places elephants have been
harried and driven from locality to locality in alleged defense of agriculture.
The records of elephants that have been killed in Malaya during the last
few years under the agriculturist's exemption are incomplete reports are
seldom sent in of elephants that have been wounded but there is reason to
believe that the Malayan elephant is on the way to extermination. It is
extremely doubtful if the yearly toll of destruction is being made up by the
yearly production of calves, and that means extinction unless a halt is called.
One must take into account the fact that wild animals when much disturbed
have a habit of curtailing their breeding, and it is almost certain that this
affects elephants as well as the other large forms of our fauna.
In Kuala Selangor, Lower Perak, Labu, and elsewhere, despite the sup-
posed sanctuary provided by Forest Reserves, the elephants have not been
left undisturbed, and have been unable to find in the areas that they have
receded to that tranquility essential to an elephant's well-being.
Where elephants have been forced to live in jungle areas which are insuffi-
cient for their normal existence, and where they have become a serious
menace to cultivation, it is advocated that they should be destroyed by per-
sons whose business it would be to undertake the work. . . .
It is an established fact that wild elephants, always providing they are not
suffering from wounds, can be driven away by fire crackers and noise. In
cases of absenteeism, which is frequently the contributory cause when ele-
phants visit native cultivation, these methods cannot be applied. A woven
ORDER PROBOSCIDEA: PROBOSCIDEANS 313
wire fence properly upkept and with a path kept reasonably clean on the
jungle boundary . . . would in most cases keep elephants from entering the
cultivated area. . . .
The removal of protection from Elephants, a measure taken in 1929, was
condemned by the vast majority of English speaking witnesses before the
Wild Life Commission of Malaya. This order was liable to accentuate the
trouble from wounded elephants and undoubtedly, as evidence showed, forced
elephants into localities where they had never been known before. This
unwise order was rescinded on the 15th of May, 1931, and the elephant
cannot now be shot at by an unlicensed person except in alleged defence of
property. . . .
Raids on native cultivation are often due to neglect. Persons familiar with
the habits of elephants can often move a herd from the vicinity of cultivation
by following them up all day until they are miles away from the locality
they visited the previous night. . . .
How do Sakai in their primitive state handle the planting of crops in ele-
phant country? Showing more wisdom than their white brothers, they leave
elephants alone. ... In the Sakai country, which lies between the main
range and the Kelantan Railway, the Sakai suffer no damage from elephants.
The elephants, not being disturbed and harried, have not learned to "answer
back."
[Some hold an opinion] that a very large percentage of the so-called damage
done by elephants is only done to patches of abandoned cultivation, and
when inhabited land is attacked it is not infrequently done by bad-tempered
elephants suffering from wounds of sorts which are caused by some home-
made bullets fired from a shot gun.
F. N. Chasen writes (in litt., May 5, 1937) : "The question of
protecting the elephant in the Malay Peninsula raises and crystal-
lizes the whole policy of local big-game preservation. Can big-game
co-exist with modern agriculturalists? My view is that the elephant
should be protected in reserves : outside the reserves he must behave
himself, or be shot. These are, of course, the extremes of the case
and a middle course is, sometimes, permissible when directed by an
experienced game-warden. The Malayan elephant is decreasing in
numbers, rapidly, in the settled areas. It is still numerous elsewhere."
Peninsular Siam. The following two accounts relate to the
uninhabited country about the northern end of the Inland Sea :
"On the plain and in the forest a herd of about 300 wild elephants
are roaming. . . . These elephants have from time to time been
captured, but their death has always resulted after some compara-
tively short time." (Havmoller, 1926, p. 365.)
"From government officials with whom I was traveling I learned
that a herd of at least 200 elephants ranges over the vast grassy
plain extending southward from near Nakon Sritamarat almost to
Singora on the west side of the Inland Sea and practically from the
Gulf of Siam to the high mountains in the west. This plain, suitable
for rice growing, is entirely uncultivated owing to the ravages of the
elephants." (H. M. Smith, 1926, pp. 365-366.)
Elephants are protected in Siam because "they are considered
314 EXTINCT AND VANISHING MAMMALS
property of the State, and therefore a special permit must be ob-
tained from the King before an elephant may be killed" (David E.
Kaufman, in litt., March 8, 1933).
Suiiiatrun Elephant
ELEPHAS MAXIMUS SUMATRANUS Temminck
Elephas Sumatranus Temminck, Coup-d'oeil Possessions Neerlandaises, vol. 2,
p. 91, 1847. ("Sumatra.")
FIGS.: Lydekker, 1916, vol. 5, p. 83, fig. 24; Pieters, 1932, p. 58, fig.
This Elephant, while still existing in considerable numbers in
Sumatra, is evidently losing ground in contact with cultivation, and
concern is felt over its future.
It is said to be characterized by its small size, its tessellated
skin, the pyriform shape of its ear, and the infolding of the posterior
edge of the ear (Lydekker, 1916, vol. 5, p. 84) .
Sumatra is the only part of the Malay Archipelago that has pos-
sessed a native stock of Elephants within historic times. Those now
found in Borneo are considered descendants of domesticated indi-
viduals introduced from the Malay Peninsula (Mjoberg, 1930,
pp. 15-16).
In 1906 W. L. Abbott (in Lyon, 1908, p. 622) saw many trails in
eastern Sumatra opposite Pulo Rupat.
Only mature males may be hunted, and the open season may not
exceed six successive months. The export of either living specimens
or the skins of Elephants is prohibited, and the export of ivory is
restricted within certain limits. During the past ten years an
average of only 350 kilograms of ivory has been exported annually
from the Netherlands Indies. The published value is only 10 to 20
guilders a kilo. Animals with very large tusks have disappeared
for the most part, and the present average weight of a pair of
tusks is estimated at 10 to 12 kilograms. Thus the above-mentioned
export figures represent the annual taking of about 35 Elephants.
(Dammerman, 1929, pp. 13-14.)
"The two principal ports to which the ivory is sent, are Singapore
and Penang. Much ivory is also carved here locally, so we may
suppose that yearly many more elephants are killed than the 35
the tusks of which are exported. With the new regulations export
of elephant-tusks weighing less than 5 kilograms a piece, is for-
bidden." (Dammerman, 1929, p. 14.)
The same author (in Skottsberg, 1934, p. 422) considers the
Sumatran Elephant threatened with extermination. According to
Pieters (1932, p. 58), the greatest danger is the encroachment of
cultivation on its habitats.
"There are still some elephants in Langkat District, but not as
ORDER PROBOSCIDEA: PROBOSCIDEANS 315
many as, say 10 years ago. There are some on Lepan, Besitang,
and Namoe Oengas. Elephants are shifting from one place to an-
other, and then come back to the starting point again." (J. Gourin.
in Hit., August 7, 1933.)
The following account is given by Heynsius-Viruly and Van
Heurn (1936, pp. 48-50) :
Very detailed reports on the elephant were received from many districts.
While some believe they will be exterminated within the next twenty-five
years, others think that they are holding their own as there are yet about
2000 elephants in Southern Sumatra alone. This estimate is, however, called
in doubt by competent observers. In the subdivision Ogan Oeloe there were at
the most about 45 elephants in 1926. There is much difference of opinion
about the damage these animals do. In Rokan they have increased so rapidly
that they have become a nuisance; nevertheless they are not hunted much.
A report from Soengi Radja relates that in 1929 a herd of 14 head was
discovered; efforts are being made to preserve them. Elephants were also
seen near Soengi Roka in May 1932. In Siak their number is estimated as
still quite large, likewise in Indragiri, although they do not appear there
in the swampy coastal districts. In the lowlands they are found only in
Reteh, and the largest herds in South Seberida in the Boekit Tiga Pdeloeh.
In the first-named district a reward of twenty-five Dutch guilders is offered
for every elephant tail. The controler of the district, which comprises the
middle course of the Siak River, paid twenty-eight such premiums in 1930.
A correspondent estimates that about 200 elephants roam over Siak and
urges the repeal of the old local regulation concerning the premiums, as
well as not extending the permits for the fire-arms kept in the kampongs.
In Djambi, Moeara Tambesi and Moeara Tebo they are fairly common;
also even now, in South west Bangko, where the controler estimates they
will be extinct within 10 years. They are very rare in Moeara Boengo. In
Djambi they are estimated at about 250-350.
There is a herd of about 30 in Korintjih, and seven in the Ophir district
(July 1932) viz: one young male, and six females. In 1915 this same group
numbered still 18. In 1916 the herd of North Korintjih were hunted by men
specially appointed for the purpose by the Demang of Korintjih and the
Civil Authority of Air Hadji. Not even the females and the young were
spared at that time. Along the Mesoedjih River elephants are caught in
pitfalls by the Natives and the younger ones are sold in Palembang. The
older ones are left to starve, in order to obtain the tusks to sell. It is generally
thought that the present regulations merely postpone the extermination of
the elephant. Only establishing extensive reserves might bring adequate
protection. . . .
The report of a herd of 14 in Soengi Radja is of much interest, for these
animals occupy a rather small area that has been completely surrounded by
cultivation for a quarter of a century, and though much hunted they have
succeeded in holding their own. The establishing 1 of a reserve here was urged
in 1929. In 1932, the Netherlands Committee for International Nature Pro-
tection requested this from the Government of the Netherlands Indies, but
as yet no actual steps have been taken.
A second important fact, emphasized by our enquiries, is that at certain
seasons elephants migrate periodically from the mountains to the lower
coastal areas. This too had been exhaustively recorded in print. It was one
of the strongest arguments used by the Netherlands Committee, when sub-
316 EXTINCT AND VANISHING MAMMALS
mitting their proposal for requesting the Indian Government to include in
the reserve certain lowland swampy areas. This has not been done at present ;
but the Committee intends calling the Government's special attention to the
new information received which further supports their claim. The protection
of the elephants remains, moreover, a separate problem. Even after re-
serves are established, and, of course, after the Decree on Hunting becomes
effective in Sumatra, the careful listing of existing herds will be imperative.
The continued gathering of data regarding each herd, in order to determine
which way they travel, their increase or decrease, and what damage they do,
will furnish the foundation for their protection and for the preservation
of the remaining herds. . . . May complete cooperation by the Department
of the Interior facilitate this task of the Netherlands Committee.
South African Bush Elephant
LOXODONTA AFRICANA AFRICAN A (Blumenbach)
Elephas ajricanus Blumenbach, Handbuch der Naturg., ed. 5, p. 125, atlas,
pi. 19, fig. C, 1797. (Selected as Orange River, South Africa.)
SYNONYMS: Elephas capensis F. Cuvier, Tableau Elem. de PHist. Nat. des
Anim., p. 149, 1798 (Orange River region, South Africa) ; Elephas ajri-
canus toxotis Lydekker, Proc. Zool. Soc. London 1907, pp. 385, 388, Aug. 1,
1907 (Addo Bush, South Africa) ; Loxodonta africana zukowskyi Strand,
Arch, fur Naturg., vol. 90, sect. A, pt. 1, p. 68, footnote, July 1924 (Kaoko-
veld, South-West Africa).
FIGS.: (Of the Tanganyika animal) M. Maxwell, 1930, 11 plates from
photographs.
Because of its great size, its strength, its tusks of ivory, its
remarkable trunk, or proboscis, and its intelligence, the African
Elephant is one of the most interesting of mammals. At first con-
fused by early naturalists with the Indian Elephant, it is, however,
so different that the two are now placed in separate genera. The
African Elephants (Loxodonta) differ in many points, such as the
structure of the tip of the proboscis, with its two instead of single
fingerlike tips, the huge ears extending back to cover the sides of the
neck as far as the shoulder blade, the forehead, which is less globular
than in the Indian species, the cheek teeth having fewer of the high
enamel prisms which form their essential grinding structure, the six
successive teeth with usually 3, 6, 7, 7, 8, and 10 prisms, respectively,
against 4, 8, 12, 12, 16, 24 in the Indian Elephant ( W. L. Sclater) .
The African Elephant is slightly the larger, but seldom exceeds
11 feet in height at the shoulder, a distance not easily measured with
accuracy even when the animal lies dead. The weight of the famous
"Jumbo" was about 6.5 tons. In color the skin is slaty gray but may
appear in life of different tints, according to the light, the dryness
of the skin, and the amount of earth clinging to it if animals have
been dusting or wallowing. A sparse coat of short stiff hairs is
insufficient to obscure the hide, but near the tip of the tail these
hairs become stout coarse bristles growing from the edges of the
ORDER PROBOSCIDEA: PROBOSCIDEANS 317
compressed terminal part, several inches in length. The upper pair
of incisors are enormously enlarged to form tusks, which are larger
in the male than in the female, or may in the latter sometimes be
lacking. These are used as weapons or in digging for roots. The
largest tusks come from Kenya Colony, with a record length on the
outside curve of 11 feet 5^ inches, and a weight for the two of
293 pounds (Roland Ward, 1935) .
The African Elephants have at various times been subdivided into
local races by systematists, but there is still much doubt as to the
value of the characters claimed, and the number of valid geographical
forms. In general one may distinguish the larger "Bush Elephants"
and the smaller "Forest Elephants," the former distinguished by
minor additional points such as the larger, more elongated ears, the
more forwardly directed tusks, less abundant hair. There is a ques-
tion whether these two types should be regarded as merely races or
as separate species, but the likelihood is that they have evolved
side by side though in different habitats, the former avoiding the
denser forests, the latter keeping more strictly to their shelter, with
the result that at present the two types seem different enough for
separation as distinct species. The larger Bush Elephants, again,
have been regarded as of several local races, of which that of South
Africa, the first to be named, is at present much reduced in numbers.
Farther to the northeast, the East African animal has been named
L. a. knochenhaueri, and the Sudanese Elephant, L. a. oxyotis. There
is still much doubt as to the validity of the characters distinguishing
these races, but until series of skulls and measurements can be com-
pared one can only await further information. The character of the
ear lobe invoked, for example by Lydekker, is so subject to modi-
fication through distortion in dried or mounted specimens that
little reliance can be placed upon it. One may then consider the
status of the Bush Elephants as a whole, with special reference to
the South African race.
In classical times elephants were found over most of Africa except
the most desert areas. There seems to be evidence that in ancient
times they were found abundantly in Abyssinia, for under the
Ptolemies, in the third century B. C., elephants for use in warfare
were captured and trained in Ethiopia on the shores of the Red Sea
and were taken thence in specially constructed boats to Egypt.
Entire army corps were sometimes engaged in their capture. In
Carthaginian days elephants were captured in Libya and in Maure-
tania among the forest-covered foothills of the Atlas Mountains.
Here, however, they have long ceased to exist and are not now found
north of the southern borders of the Sahara. In the eastern Sudan
elephants still occur in small numbers (I myself saw their "sign"
on the Blue Nile, near the Abyssinian border in 1913), but have
318 EXTINCT AND* VANISHING MAMMALS
long ago retreated from the borders of the Red Sea. Swayne wrote
a quarter of a century or so ago: "There is practically no elephant
shooting to be got in Somaliland north of the Haud Plateau, or in
the Haud, at the present time. In the gorges which descend from
the highlands of Abyssinia to Ogaden in the country about the
head-waters of the Webbi Shabeyleh and Juba Rivers there are
still plenty of elephant. A few herds, it is believed, wander down
those river valleys to the Marehan Country far to the south-east
of Berbera." A certain amount of ivory-hunting by natives may
keep these herds in check. But recent travelers up the Nile report
large numbers of elephants in the practically impenetrable papyrus
swamps of "the Sudd" where they will doubtless find sanctuary for
a long period to come. Between this area and Uganda there are
large numbers of elephants, and in the Kenya forests and thorn-
bush a good many still survive.
For the purposes of the present report, chief interest centers in the
elephant of South Africa, which nowadays with increasing settle-
ment of this part of the continent comes into close association with
white men and has had to suffer in consequence. The following brief
notes are given in summary from Shortridge (1934, vol. 1, p. 362) and
W. L. Sclater (1900) . This, the typical race of African Elephant, at
present seems to be characterized in part by its rather short stout
tusks as compared with the other Bush Elephants, but how far this
may have been due to the process of selecting largest tusks and elimi-
nating these animals in ivory hunting is not clear. "In the days of
van Riebeck (1653) elephant were plentiful as far south as the Cape
Peninsula," but by the beginning of the next century seem to have
become rare, for according to Theal the last one shot in this region
was killed "just beyond Cape Flats in 1702; the expedition of Cap-
tain Hop, in 1761, found plenty just north of the Oliphant River
in what is now the district of Clanwilliam, while in the eastern half
of the Colony, elephant hunting was regularly pursued till about
1830. ... In Natal a few survived till 1860; in the north the
hunters of the early part of the century made large bags near
Kuruman ; Harris in 1836 shot chiefly in Magaliesberg of the western
Transvaal; Gordon Gumming in 1846 in Sechele's country in
northern Bechuanaland, and Livingstone and Baldwin, in 1849 and
1858, found elephants innumerable on the Botletli River and near
Lake Ngami, and finally Selous' hunting ground in the seventies
and early eighties was in what is now Matabeleland and Mashona-
land." Elephants were formerly so plentiful in the southeastern part
of the Cape that an important ivory market was established in 1824
at Fort Willshire. After 1860, however, the herds in the Knysna
Forest and the Addo Bush were placed under government protection.
The last elephant in Zululand was said to have been a solitary bull,
ORDER PROBOSCIDEA: PROBOSCIDEANS 319
which was killed in February, 1916, and its skeleton is now mounted
in the Natal Museum. In Matabeleland, elephants existed in large
numbers in 1872 and had been little hunted, but in the few succeeding
years Lo Bengula's hunters in addition to Europeans swarmed into
the region and in three years took out an estimated 100,000 pounds
of ivory. Even then, tusks over 70 pounds in weight were rare, and
the average was 40 to 50 pounds, rather small as compared with
those farther north. By 1902 elephants had disappeared from the
Transvaal, but in late years a few have come back into Kruger Park
from adjacent areas of Portuguese East Africa. Shortridge sums
up the present situation in the Cape Province and adjacent terri-
tory: "Scattered and comparatively small herds of elephant still
wander in Ngamiland, Southern Rhodesia, Portuguese East Africa.
... In the Cape Province, the remnant of a herd is preserved in
the Addo Bush. There may still be half a dozen or so in the Knysna
Forest." In South-West Africa, there may be from 600 to 1,000 head
in the Kaokoveld, but larger estimates are probably unwarranted.
The other region where they occur is in the Caprivi, where two fair-
sized herds are said to survive, one near the Kwando River, the
other between Popa and Kagera. There are still elephants in south-
western Angola and especially along the Kwando in the southeastern
part. An estimate of the elephant population of any district is not
easy to make, however, since on account of their wandering habits
the same animals may appear within a short time at points far apart.
In South Africa, aside from the restricted herds of Caprivi and the
Kaokoveld, there exist four other herds: (1) that in the Knysna
Forest, said to number, in 1935 about a dozen animals, which are
under Government protection; (2) the Addo Bush herd, near Port
Elizabeth, numbering, in 1933, about 16; (3) the Kruger Park herd,
which seems most favorably situated and is believed to receive
occasional increments from animals seeking this sanctuary from
adjacent Portuguese territory; and (4) a small number that occa-
sionally appear in straggling parties from across the Limpopo in
times of drought and enter the northern Transvaal. Concerning
the Addo Bush herd, in the early part of 1920 its numbers were 126,
more than could well be maintained there, and so by Government
order 110 were killed, and the remnant was confined to a more limited
space, which apparently the animals more or less recognize and keep
within its limits. A boring to supply them with water has been made
to help in keeping them within these bounds, but the difficulty of
restricting their wanderings is not easily overcome. In Kruger
National Park there are said to be (1933) approximately 150-200
elephants, in five separate groups. They tend to spread out from
their fastnesses among the reed beds of the Letaba River (1934) .
Apart from its great interest, from both esthetic and zoological
320 EXTINCT AND VANISHING MAMMALS
points of view, the African Elephant has for centuries supplied a
large part of the world's demand for ivory; it is a source of meat
for many native tribes; and in recent years it has again been the
subject of attempts at domestication, in this instance in the Belgian
Congo, where imported mahouts from India as well as tame Indian
Elephants have succeeded in rendering the Forest Elephant more or
less tractable. Their timidity, however, often impairs their useful-
ness, while the large amounts of food they require add to the diffi-
culty of an economical value. According to Lavauden (1933, pp.
21-22), in 1921, ivory to the amount of 800 tons of elephants' tusks
was sent to the world's markets; in 1925 this had fallen to 500 tons,
but the average weight of the tusks had considerably decreased as
well. Elephants, on the other hand, often do much damage to the
crops of the agricultural natives, wrecking their fields, granaries, and
even houses at times. This damage, although "it is very doubtful if
it would amount to 1 percent of the entire crop," is nevertheless at
times a considerable loss, and of late years measures have been taken
in countries under British rule to cope with this, by appointing an
official to undertake elephant control through killing a certain num-
ber in areas where they are reported to be doing such damage. In
his book Elephant, David E. Blunt (1933), who had charge of this
work in East Africa, reports that elephants seem very quickly to
learn the bounds of regions to which they must be confined, and
after a few of a marauding herd are shot the trouble to plantations
is stopped for at least the time being. Thus while it is possible by
this means to reduce greatly the elephant damage in agricultural
areas near large forests or other country inhabited by herds of these
animals, it is likely that with increase of settlement this protection
of crops will become less needed, and the animals will gradually give
way. Nevertheless there will undoubtedly be plenty of elephants in
some sections of Africa for many years to come, in spite of hunting.
Moreover, these will prove an asset on account of the returns from
purchases of big-game licenses and additional fees for each elephant.
In East Africa animals with tusks under fifty pounds in weight (the
two together) may not legally be killed under penalty and confis-
cation of the ivory. This limit, according to Brocklehurst (1933),
has been lately reduced in Abyssinia from 30 to 20 pounds so that
females now are killed.
It appears from statistics that Uganda is likely to be one of the
regions where elephants will long hold out and may be an asset in
the way mentioned. In 1929, the Game Department reported a kill
of 1,439 elephants, of which 1,135 were accounted for by the Govern-
ment control operations. In 1931 the Game staff killed 1,211; in
1933, the number was 1,380, and yet "with the exception of the Toro
district, the southern portion of West Nile, and possibly the
Mubende district, there is no reason to believe that elephant num-
ORDER PROBOSCIDEA: PROBOSCIDEANS 321
bers are other than steadily increasing in all parts of the protec-
torate in which this great beast occurs." In 1934, it is said that
2,716 elephants were killed in Tanganyika Territory. Taylor, in
East Africa, July 9, 1936, believes that the method of control adopted
is "the most humane method possible of enabling men and elephant
to live in peace and concord in one territory." With the stopping
of such methods of slaughter as once were practiced by natives in
encircling elephant herds by grass fires and using pitfalls, and with
the reduction of poaching for ivory, and the licensing of hunters, no
doubt the hazards for the species are sufficiently lessened to counter-
balance the large numbers just noted that are killed in control
measures. The ivory from such elephants as are killed in this way
is Government property and a source of revenue. Ivory is also a
regular product of the Belgian Congo, where many animals must
annually be killed, although at the present time this requires special
license.
For the future, the opinion of those conversant with the situation
seems to be that in South Africa the relatively small areas of national
reserves may continue to hold elephant herds indefinitely, but the
size of the herds must be regulated by the area of the reserve and
its suitability to their needs. With reduction to small numbers there
is always a danger of an unlooked-for change which may be un-
favorable. In South-West Africa, the numbers yet remaining are
under government protection, so far as it may be enforceable, but
the elephants here doubtless owe their continuation quite as much
to the inaccessibility of their habitat. In the less settled parts of
East Africa, they will continue in numbers and with the present
efficient supervision of the game departments should prove on the
whole a decided asset and attraction, notwithstanding a certain
amount of local damage to plantations. In Uganda, where the herds
are still abundant, there is evidence of slight increase in numbers in
some districts, while in the great papyrus swamps of "the Sudd" of
the upper Nile, they are present in great numbers and are likely to
find this a safe retreat. Airplane photographs taken by the late Mar-
tin Johnson in this region show some astonishingly large herds. The
game warden of Uganda in his report for 1925 believes that with
the spread of settlement and development elephants will have to be
killed out or "expelled" from certain areas, but that, since extermi-
nation is impossible and impracticable, good sanctuaries are neces-
sary, which shall protect the main breeding areas of the herds.
Elephants quickly learn to recognize the areas in which they are
free from molestation, so that this trait will help to keep them within
such bounds. A proper sanctuary, however, must include sheltered
valleys with abundance of food and sufficient water, else at periods
of drought the animals will move off in search of better localities.
G. M. A.
322 EXTINCT AND VANISHING MAMMALS
Order PERISSODACTYLA: Odd-toed Ungulates
Family EQUIDAE: Horses, Zebras, and Asses
Some conservative zoologists recognize but one genus in this
family, while granting subgeneric status to the Horses (Equus),
Zebras (Dolichohippus, Hippotigris) , and Asses (Asinus) . Others
raise these subgenera to generic rank, and Shortridge (1934, vol. 1,
p. 397) proposes an additional genus (Quagga) for the Quagga and
Burchell's Zebra. Dr. Allen maintains a conservative viewpoint and
employs the generic name Equus for all the Zebras (including the
Quagga), while I prefer to keep both the African and the Asiatic
Asses in a separate genus, Asinus. The single surviving species of
Wild Horse (Equus przewalskii) is now confined to Mongolia. The
nine forms of Zebras (two extinct) occupy eastern and southern
Africa. One extinct and two living forms of African Wild Asses
(Asinus atlanticus and A. africanus subspp.), with ranges in the
northern and eastern portions of that continent, are herein recog-
nized; also six forms of Asiatic Wild Asses (Asinus hemionus) ,
ranging from Mongolia and Tibet to Syria. The generally pre-
carious status of the family is indicated by the fact that all but
one of the Asiatic forms and all but six of the African forms are
treated in the following pages.
Przewalski's Horse; Mongolian Wild Horse; Mongolian Tarpan
EQUUS PRZEWALSKII Poliakov
Equus Przewalskii Poliakov, Izviestiia Imper. Russk. Geogr. Obshchestvo,
vol. 17, p. 1, 1881. (The type specimen was obtained by a "hunting-
expedition sent by M. Tihonof from the post Zaisan to the sand deserts
of Central Asia" (Poliakof, 1881, p. 19). Type locality restricted by
Harper (1940, p. 195) to the oasis of Gashim, eastern Dzungaria (approxi-
mately lat. 44 30' N., long. 90 E.).)
SYNONYM: Equus hagenbecki Matschie (1903).
Fios.: Poliakov, op. cit., pi. 1; Przewalski, 1883, pi. facing p. 40; Lydekker,
1901, p. 284, fig. 65; Salensky, 1902, pi. 1, pp. 12, 16, 17, figs. 2-4; Proc.
Zool. Soc. London 1902, vol. 1, pi. 13; Matschie, 1903, p. 582, fig.;
Ridgeway, 1905, p. 27, fig. 18, p. 29, fig. 19; Wrangel, 1908, vol. 1, p. 3,
fig. 1; Lydekker, 1916, vol. 5, p. 8, fig. 4; Peake, 1933, pi. 31, fig. a;
Pocock, 1937, p. 715, fig.; Reed and Lucas, 1937, p. 129, fig. 44; Schmidt,
1938, pi. 10.
A very special interest attaches to this animal, as the only truly
wild horse surviving in the world today. There is a remarkable
dearth of first-hand information concerning it, especially during
the past quarter of a century or so. Only one of the numerous
scientific expeditions to Central Asia during recent years seems to
have come into contact with it. It is somewhat doubtful if the
alleged Mongolian Tarpans now exhibited in American zoos are
purebred animals.
ORDER PERISSODACTYLA: ODD-TOED UNGULATES 323
Przewalski's Horse is distinguished from other horses by its erect
mane and lack of a forelock. The following description is derived
from Salensky (1902, pp. 7-18), who had more than a dozen speci-
mens at his disposal, rather than from Poliakof (1881), who had
only one.
This species is of the size of a small ordinary horse; grown male
with a height at the rump of 1,240 mm.; head relatively larger than
that of the Wild Ass ; average length of ears, 140 mm. ; mane erect,
highest (160-200 mm.) in the middle of its length; tail long, reach-
ing in some individuals nearly to the hoofs, and provided on the
dorsal side toward the base with short hairs, elsewhere with long
hairs; hoofs rounder than those of the Kiang and the Kulan;
"chestnuts" on all four limbs.
Winter pelage lighter than that of summer; yellowish on the back,
becoming lighter on the sides and almost white on the under parts.
Summer pelage much shorter than that of winter, smooth, not wavy;
back and sides light reddish brown, gradually changing to yellowish
white on the belly; head colored like the back, but white on the
muzzle about the nostrils and on the lips; ears light brown basally,
darker at the tips; inner surface of ears white. Pronounced tufts of
hair on sides of head in winter, and along entire lower part of head
in summer. Mane dark brown, with shorter tufts of light gray hairs
on each side; a median dorsal stripe of reddish brown, about 5 mm.
wide, and distinctly visible only in summer pelage, extending along
the entire back and on to the tail; a brown or black shoulder stripe,
more noticeable in summer than in winter ; lower part of limbs more
or less black (occasionally gray in younger animals) ; a black ring,
up to 80 mm. wide, bordering the hoofs; inner side of legs gray,
generally with distinct bars, up to the knees.
The principal range seems to have been on both sides of the Altai
Mountains in western Mongolia and in Dzungaria. But Prejevalsky
(1876, vol. 2, p. 170) also reported Wild Horses much farther south,
in western Koko Nor and in southeastern Chinese Turkestan : "Wild
horses, called by the Mongols dzerlik-adu, are rare in Western
Tsaidam, but more numerous near Lob-nor. They are generally in
large herds, very shy, and when frightened continue their flight for
days, not returning to the same place for a year or two. Their colour
is uniformly bay, with black tails and long manes hanging down to
the ground. [This last expression is, of course, wholly erroneous as
applied to the manes.] They are never hunted, owing to the diffi-
culties of the chase." This report, apparently based upon native
information, does not seem to have been substantiated by later
records, and is open to question. In this connection, however, it may
be recalled that Sven Hedin has remarked (1903, vol. 1, p. 357) on
324 EXTINCT AND VANISHING MAMMALS
the increasing scarcity of even the Wild Camel east of Lob-nor,
owing perhaps to the increasing desiccation of the region.
The animal must have been somewhat rare in the Altai region
even before the advent of Europeans. Atkinson (1858), who made
extensive explorations in that region about the middle of the past
century, and comments frequently on the other large animals, does
not refer to any personal encounter with the Wild Horse. However,
in a later work (I860, p. 325), he describes the Kirghiz method of
hunting "wild horses, which at this season [May] are found in
great herds near the foot of the mountains" beyond the Hi River,
apparently toward Issyk Kul. But his description of these horses
as "varying in colour from black, bay, grey and white" creates
considerable doubt as to whether they were truly wild or merely
feral. Possibly the herds were composed of a mixture of both kinds
of animals. Atkinson gives the Kirghiz name for the wild horse
as "muss."
Brehm (1876, p. 339) received a report of a second kind of Wild
Horse (besides the Kulan), called "Surtake," which was said to
occur about 250 versts southeast of the boundary post of Zaisan, in
the Kanabo area. It was described as light yellow, with many light
spots and a shorter tail than the Kulan's.
Younghusband, referring to the region about the southern base of
the eastern Altai, at about long. 96-100E., says (1888, p. 495) :
"We . . . saw here . . . wild horses too the Equus Prejevalskii
roaming about these great open plains."
Ten years after Przewalski's discovery, the brothers Grum-Grshi-
mailo took some specimens in 1889 at the oasis of Gashun, northeast
of Guchen in eastern Dzungaria (Wrangel, 1908, vol. 1, p. 2) .
The following report of Grum-Grshimailo (in Morgan, 1891, pp.
217-218) probably refers chiefly to the Gashun area: "Springs
enable the numerous animals inhabiting Dzungaria to exist; of
these the most interesting is Prejevalsky's horse. . . . Prejevalsky
himself, though he crossed the desert of Dzungaria in three several
directions, never came across any of these wild horses, and if he
wrote otherwise he was mistaking kulans he had seen in the distance
for wild horses, a mistake the most experienced hunters are liable
to make, for at that distance it is almost impossible to distinguish
between them. . . . We were the first Europeans who, for twenty
days, made a study of these interesting animals, adding the skins
of three handsome stallions and one mare to our collection."
Salensky (1902, pp. 2-3) records specimens from the following
localities, chiefly in or near the Dzungarian Gobi: Gashun; the
Kobdo region; behind the Baitik-Bogdo (Charamelechetai) ; between
Nursu and Simigendse; Ebi Spring, near the Kobdo-Barkul route;
Guchen Lake; and the River Bulunga. He gives the range (p. 63) as
ORDER PERISSODACTYLA I ODD-TOED UNGULATES 325
extending north to the Urungu River and Kobdo ; east to longitude
90-91 E.; south to latitude 46 N.; and west to longitude 84 E.
[=86?]. (This range is too restricted on the east and south.)
In 1899 three newborn foals were captured and in the following
year were brought to the estate of Herr Falz-Fein in Ascania Nova,
southern Russia (Salensky, 1902, p. 20) .
In 1901 Carl Hagenbeck sent a large expedition to Dzungaria for
Wild Horses. His animals were caught in three different districts
lying south of the Mongolian city of Kobdo. In the west the area
consists of a wide plain, bordered on the east by the Altai Mountains.
It is bordered on the north by the Kui-Kuius River, and on the
south by the Urungu River, both of which rise in the Altai and
discharge into the Tusgul [Ulungur?] Lake. This lake forms the
western boundary of the plain. The second area is a plain which
lies about 322 km. south of Kobdo and is enclosed by the Altai
Mountains. The third group comes from the vicinity of Zagan Nor
[apparently near long. 95 E.]. Foals from the three groups differ
in color characters, though quite alike in general appearance
(Wrangel, 1908, vol. 1, pp. 2-3) .
The foals are dropped between the end of April and May 20. Their
capture takes place as follows. Hundred of Mongols lie in ambush
behind hills. As soon as they see a considerable number of mares
and foals together, they rush upon them with loud cries. Since the
foals can not keep up with the fleeing mares, the Mongols soon catch
them with nooses on long poles. They are then conducted to near-by
corrals, where Mongolian mares are ready to take over the duties
of foster-mothers. Of the animals thus captured by the Hagen-
beck expedition, 28 arrived in Hamburg in 1901 (Wrangel, 1908,
vol. 1, p. 4) .
"There is no doubt that the wild-horse . . . also inhabits the
northern portions of that region [ Dzungaria]. We were never lucky
enough to see any, but the natives, both Kalmuk and Kazak, all
told the same tale, often volunteering the information that, in
addition to the kulon, there were wild-horses. . . . They said, the
meat was not so good [as the kulon 's]. They told us that there
were large herds of them in the vicinity of Lake Ulungur, and east-
wards along the southern foot of the Altai; also north of that range."
(J. H. Miller, in Carruthers, 1913, p. 608.)
From a point on the north side of the Altai, about 100 miles west
of Ikhe Bogdo, R. C. Andrews reports (1926, p. 322) : "The wild
. . . horses were two hundred miles to the southwest, they [some
Chinese caravan men] said, just above the border of Chinese
Turkestan."
Morden writes (1927, p. 286) concerning a place in eastern Dzun-
garia, northeast of Kucheng: "Around the spring, which our men
326
EXTINCT AND VANISHING MAMMALS
we heard that the wild horses of
. . were sometimes seen by cara-
said was called Kainar Bulak . .
western Mongolia and Dzungaria
vans approaching the place."
The range of the species was extended eastward by Lattimorc
(1929, p. 228). In a journey of 1926, he mentions passing the "Yeh-
ma Ching" or Wild Horse Well, which is situated in the Khara Gobi
FIG. 33. Mongolian Wild Horse (Equus przewalskii) . After photograph
in Brehm.
west of Edsin Gol, at about latitude 42 N., longitude 98 E. "They
say that on this fringe of the Khara Gobi there are wild horses
(equ,us prjevalskii] and wild asses." This recalls Ridgeway's state-
ment (1905, p. 28) : "Mr. Hagenbeck informs me that wild horses
of another variety are said to exist 600 miles south of Kobdo.
that is, somewhere in the great Gobi desert."
Teichman also passed by the Wild Horse Well, in 1935, and makes
the following remarks (1937, pp. 74-75) : "The Hardt-Citroen expe-
dition followed from Suchou to Mingshui a camel trail which took
them through this range. They found a region of rich pastures,
abundant water, the haunt of ... the wild horse and wild ass.
. . . Wild horses and wild camels are said to exist in this neighbour-
ORDER PERISSODACTYLA : ODD-TOED UNGULATES 327
hood. We saw no direct evidence of either. The wild ass is common
and is often, with characteristic Chinese lack of accuracy, referred
to as Yeh Ma ('wild horse') , which may explain the name of Yeh-ma
Ching."
Reymond, the zoologist of the expedition mentioned by Teichman,
records (1932, pp. 807-809) a Wild Horse seen in May, 1931, and a
carcass recently devoured by Wolves near the northern border of In-
ner Mongolia at longitude 105 30' E., latitude "40" [ =42] N. The
skull of the latter was identified by Professor E. Bourdelle of the
Paris Museum. Reymond also heard that a favorite haunt of Wild
Horses was the plateau of Pei Chan, occupying the extreme western
triangle of Inner Mongolia. Other members of the Haardt-Audouin-
Dubreuil Expedition (as it is here called) saw in 1931 two solitary
animals in this general region: one in June, 20 km. west of Hou
Hung Chuan (long. 96 E., lat. "40" [ = 42] N.), and one in
December near Hsin Hsin Chia on the Kansu-Sinkiang frontier.
The first of Reymond 's records is by far the easternmost one to date.
"Przewalski's wild horse is found in small herds in Chinese Tur-
kestan (Sinkiang) and Western Mongolia. It does not appear to be
at all numerous, and should be protected if possible, if only because
it is the sole surviving true wild horse in the world to-day. It is too
small to be of any economic value, the so-called Mongol pony used
by the Mongols and other Central Asian people easily supplanting
it. The latter is probably a cross between Przewalski's horse and
various domestic breeds, and is sufficiently hardy to live in the great
wastes of Central Asia in a feral state." (Sowerby, 1937, p. 250.)
Antonius writes (1938, pp. 558-559) :
The statements of the brothers Grum-Grshimajlo and the expeditions of
Falz-Fein and Hagenbeck for obtaining living specimens make it possible
to give the geographic distribution about 1900. There was only one district on
the Northern ranges of the Ektag Altai: the neighbourhood of the Zedsig-
Nor, called also Zagan-Nor, and three or four in the deserts on the southern
ranges: the steppes on the upper Urungu, the Ebi-mountain, the Gashium-
desert. If my Russian information is correct and I have, alas! no doubt
that it is, the Przevalski horse has been extirpated since the great war and
the Russian revolution, the old fork-muskets of the Mongolian hunters
having been replaced by modern fire-arms of great power. Therefore it is
probable that the descendants of the Falz-Fein and Hagenbeck-imports living
in Ascania Nova, Woburn Abbey, and in a few Zoos in Europe, America, and
Australia, are the last survivors of the Przevalski-horse, and of the true wild
horse in general.
There would appear to be considerable likelihood that Przewalski's
Horse, if not exterminated outright, has proceeded far along the road
to extinction through dilution with the ponies of the Mongols. In
Salensky (1902, p. 21) we find a report of domestic mares mating
with wild stallions. In remarking on the variations in color exhibited
328 EXTINCT AND VANISHING MAMMALS
by Przewalski's Horse in different parts of its range, Lydekker says
(1912, p. 89) : 'These differences suggest that there has been some
admixture with domesticated breeds." Who can even say that the
type specimen was purebred?
The chances of mixture with domestic stock are suggested by
Carruther's report (1913, pp. 532-533) of "immense droves of horses
running half wild over the prairies" in the vicinity of Barkul,
southern Dzungaria. "We . . . believe that the real 'wild animals' of
the Barkul basin signify the great herds of unridable horses which
roam untamed over the steppes. These form an Imperial Stud, and
are said to number fifteen thousand, the pick of which are trans-
ported yearly to Pekin."
Domestication. On this subject Peake (1933, pp. 99-100) says:
There can be no doubt that the horse was first tamed in the grasslands
of Central Asia, for it is only there that the wild horse, known as Przewal-
sky's horse, is to be found to-day. The first mention of the horse that has
been met with is in a document from Babylon, dating from before 2000 B. C.,
in which it is called the ass from the East. This indicates the direction from
which it came, but it does not seem to have been introduced into Mesopotamia
before the arrival of the Kassite conquerors about 1746 B. C. We have,
however, some reasons for believing that it had been tamed at an earlier
date. Into the north of Mesopotamia there had arrived some centuries earlier
a people known as the Kharians, some of whom were later called the Mitanni.
These, we know, were great horsemen. They occupied the country around
Haran, which lies between the Tigris and the Euphrates just below the points
at which they emerge from the mountains, and they seem to have arrived in
that district from the North-east, probably from the Persian plateau, whence
later the Kassites descended upon Mesopotamia. The horse was well known
also to the Hittites, the capital of whose kingdom lay in the centre of Asia
Minor. These people are believed to have arrived there about 1900 B. C.
from the North-west. All this evidence tends to show that the horse was
used as a means of transport both in Persia and upon the Russian steppe
well before 2000 B. C. It seems likely that it was first tamed in that part
of the world, or still farther east in Mongolia, as early as 3000 B.C., if not
before that date.
Lamm Wild Horse
EQUUS sp.
Surprising news of a generally overlooked and probably extinct
Wild Horse in the Kolyma Basin of northeastern Siberia is fur-
nished by Pfizenmayer (1939, pp. 112-113). While excavating the
frozen carcass of a Mammoth on the Beresovka River in 1901, he
questioned two Lamut visitors as to
what sort of wild animals they found in their distant hunting-ground on the
Omolon. To our great astonishment Taitshin mentioned the wild horse. As
zoologists thought wild horses existed only on the steppes of central Asia, we
received his statement very doubtfully, though Amuksan confirmed it by a
quite professional imitation of horses neighing. The reliability of the natives
ORDER PERISSODACTYLA I ODD-TOED UNGULATES 329
is such that we did not imagine they were spinning a yarn when they told us
about the wild horse in the tundras bordering the forests of this vast area.
They described in detail its size equal to that of a Yakut horse its long
whitish -grey hair, and its flesh, which was very fat and pleasant to taste.
If the description were really that of wild [= feral] horses, it was a puzzle
how and when their tame ancestors could have reached this quite uninhabited
Arctic region. And if they actually existed in the district between the two
largest tributaries of the Kolyma the Omolon and the Anjui which had
never yet been explored by any scientist, it was a very interesting matter
which scientists would find it well worth while to investigate.
Pfizenmayer's assumption that these horses were descended from
tame ancestors is by no means necessarily correct. In this connection
it is of interest to recall Hay's opinion (1913, p. 9) that in the Yukon
Basin and adjacent parts of Alaska horses "became extinct about
the middle of the glacial epoch."
Pfizenmayer writes further (pp. 176-177) :
The prehistoric wild horse to which is probably related the animal that
Przevalski, the Russian explorer of Asia, discovered in 1870 and called a
wild horse has left remains everywhere in central and northern Siberia.
There is hardly one place on the banks of rivers and lakes in the district
of Yakutsk in which prehistoric remains of animals have been found that
has not yielded skeletal fragments of the prehistoric wild horse.
In the landslide on the Beresovka we found, among the debris between
the larch trunks lying around in confusion and the masses of fallen earth, the
perfectly preserved upper skull of a prehistoric horse, to which fragments of
muscular fibre still adhered. . . .
An exiled student told me, in Verkhoyansk on my way back from Kolymsk,
that an ivory hunter had found the carcass of a horse four years before,
sticking half out of the frozen earth in a fissure in the bank of a lake in the
tundra, in the northern part of the district. According to the description
by the man, who puzzled over the find unusual there the parts of the
body sticking out of the ground showed a covering of very long greyish-white
hair. Certainly the Yakut horse, a vigorous breed of pony, with probably a
strain of the wild horse, also has long hair to protect it from cold. Since,
however, there were no Yakut settlements for hundreds of miles round the
site of the find, we may conclude that the body was that of a prehistoric
wild horse. But it was naturally neither investigated nor salvaged.
Determination of the relationship between this prehistoric horse
and the Recent Lamut Wild Horse must await the acquisition of
suitable museum material.
In commenting upon some earlier publication of Pfizenmayer's
findings, Antonius writes (1938, p. 559) : "One might suppose that
these white horses are descendants of any semiferal Jakute-breed
the Jakutes being the most northern horse-breeders, but ... it,
could be possible that these wild horses of the Lamutes are the last
survivors of a northern branch of the Caballus-Group, and there
are some indications for a formerly much greater distribution includ-
ing not only Eastern Siberia, but also Alaska. Since the excavation
of the Beresovka-Mammoth there are no records of the Lamute
horses."
330 EXTINCT AND VANISHING MAMMALS
The plausibility of a white horse on the Siberian tundra is en-
hanced by Janikowski's account of present-day domesticated descen-
dants of the Wild Horse of Poland. He refers (1942, p. 682, figs. 4-5)
to two survivors "which had the remarkable and unique property of
turning white in winter .... Every winter they changed the
mouse-grey summer coat . . . into a snow-white coat, only the face,
fetlocks, mane and tail retaining the dark colour."
European Wild Horses
EQUUS spp.
The taxonomic and nomenclatural status of European Wild Horses,
especially during the more recent historical times, becomes extraor-
dinarily complicated owing, on the one hand, to the lack of ade-
quate material and authoritative data and, on the other hand, to
the probability of interbreeding with domestic types. The technical
nomenclature is too involved to be discussed in detail in this brief
account. Opinions differ as to whether some of the described forms
were truly wild or were mixed with the blood of domestic horses.
Only purebred wild animals come properly within the scope of the
present report.
Remains of Pleistocene or older horses have been recorded in
various localities from India and Turkestan to Spain, France, and
England ; some of these were doubtless ancestral to the present-day
horses.
In classical times Strabo reported Wild Horses in Spain. In the
Middle Ages there are records of Wild Horses in Germany, Poland,
Lithuania, and Russia; but there is some question as whether all
of these records refer to purebred wild animals.
In 1768 S. G. Gmelin collected four Tarpans in the Government
of Voronesh, Russia. Pallas (1811) reported Wild Horses as inhab-
iting the steppe country from the Dnieper to the Altai and beyond
into Central Asia, but as partly mixed with feral animals. Hamilton
Smith (1845-1846, pp. 160-166, pi. 3) received information from
Cossacks and others early in that century concerning truly wild
animals in Russian Turkestan and Mongolia.
Antonius (1912, p. 513) mentions three animals captured alive in
Russia as late as the period 1853-66 ; he considers these the last Wild
Horses taken in Europe. However, Lydekker (1912, p. 81) suspects
that even Gmelin's specimens were hybrid Tarpans, and it is all
the more to be doubted that the animals of 1853-66 were purebred.
Antonius (1912, p. 516) has given the name of Equus gmelini to the
three last-mentioned animals, at the same time stating that Gmelin's
specimens were probably though not certainly identical with them.
If the specimens on which the name Equus gmelini was based were
ORDER PERISSODACTYLA : ODD-TOED UNGULATES 331
not purebred, the name can hardly be applied to their truly wild
forebears in Russia, which have been extinct for probably more than
a century. In the present rather chaotic state of the nomenclature,
I feel unable to fix upon any one of the numerous names proposed as
applying strictly and validly to the form represented by the last
truly wild and purebred horses of Europe.
A few quotations from the literature will indicate some of the
varying opinions on a complex subject. The later accounts can
hardly refer to purebred wild animals. Poliakof (1881, p. 20) says:
The information regarding the tarpan collected by Rytchkof, Gmelin,
Georgi, and Pallas is of so contradictory and confusing a nature that many
zoologists have decided that the so-called wild horses, or "tarpans," were
not, strictly speaking, wild, but tamed horses which had resumed their
wild state on recovering their liberty .... Pallas . . . assumed the feral horses
. . . roaming over the steppes of the Yaik [Ural] and the Don as well as on
that of Baraba to have originated from domesticated horses owned by Kirghiz,
Kalmuks, or other wandering tribes, and to have become wild. . . . Un-
fortunately we have no reliable information on this legendary tarpan since
the end of the last century, not a single traveller either in Siberia or Russia
having communicated any information concerning it during the present
century.
"The nearest approach to truly wild horses existing at present are
the so-called Tarpans, which occur in the steppe-country north of
the Sea of Azoff, between the river Dnieper and the Caspian. They
are described as being of small size, dun color, with short mane and
rounded, obtuse nose. There is no evidence to prove whether they
are really wild ... or feral." (Flower, 1892, p. 83.)
Calinescu (1931, p. 82) reports Equus caballus gmelini as sur-
viving in Moldavia, Rumania, up to 1716.
Vetulani (1933, pp. 281-282) gives the following account for
Poland. Hacquet (1794) describes wild horses kept in a zoological
garden near Samosch. They increased to such an extent that some
were shot and others were sent to Lemberg for use in combats with
carnivores. In Kajetan Kozmian's reminiscences of the years 1780-
1815 (published in 1858), we read likewise of wild horses in a
zoological garden near Zamosc or Samosch. They were allowed to
become extinct, apparently because in winter it was necessary to
provide barns and hay for them. These two references concern the
last wild Forest Tarpans of Poland ("E. c. gmelini ssp. silvatica") .
From Brincken (1874) we learn that this stock was derived from
the last wild horses in the Forest of Bialowies, and that finally they
were captured in the zoological garden near Samosch and divided
among the peasants. This represented the last stage in the domesti-
cation of the European Wild Tarpan. We still find in this vicinity
representatives of the Forest Tarpan type in an especially pure and
332 EXTINCT AND VANISHING MAMMALS
typical form. It is proposed to introduce and preserve some of them
in the Polish National Park at Bialowies.
"According to Vetulani, the enigmatic wild white horse described
by Herodotus as grazing in the northern marshy land may well have
been the Polish wild pony grazing in the Polesie bogs situated close
to the Bielowieza Forest" ( Janikowski, 1942, p. 682) .
According to Niezabitowski (1934, p. 196), E. gmelini Antonius
lived formerly in the steppe region of eastern Poland, while E.
gmelini silvaticus Vetulani inhabited the Bialowies Forest up to
the middle of the eighteenth century.
Heptner reports (1934, pp. 431-433) that the last example of
Tarpan was seen in 1914-18. It lived at that time on an estate in
Dubrowka, Mirgorod district, Government of Poltava, and was
very old. It had been purchased as a young animal from German
colonists, who shortly before had destroyed a small herd of wild
horses. Hitherto the last Tarpans in South Russia were supposed to
have died out in the 1870's. They survived longest on the steppes
of the Government of Cherson.
From the foregoing it may be gathered that it is virtually impos-
sible to state even approximately when the last truly wild repre-
sentatives of the genus Equus perished in the various European
countries. Even the names that should be applied to them are far
from settled. The type of Equus caballus caballus Linnaeus is the
Scandinavian domesticated horse of the time of Linnaeus obviously
at least subspecifically distinct from the Russian Wild Horse.
Certainly all Wild Horses are now extinct, with the exception of
Przewalski's Horse of Central Asia.
For a fuller account of the Wild Tarpan and its relations, Lydekker
(1912, pp. 71-116) may be consulted. For a discussion of some of the
nomenclatural problems involved, see Harper (1940, pp. 195-197).
Antonius (1938, pp. 557-558) gives the following illuminating
account of the caballus-group of horses :
In times not long before the beginning of historical days there were true
wild horses of the Caballus type spread over the whole Eurasiatic continent
from the Atlantic to the Pacific, and from the shores of Northern Siberia
to the Indian Ocean. Only two, or at the utmost three, of the many local
and geological races have survived until our days. The first of these was a
mouse-dun horse, which Albertus Magnus, the great interpreter of Aristotle,
means when he calls the colour of the wild horse "cinereus," i. e., ash-coloured.
There can be little doubt that these ash-dun or mouse-dun wild horses were
often intermingled with escaped domestic horses of the feral breeds, thence
spread over Europe. But there are some indications by which in many cases
their true wild nature may be ascertained. The one is the high value of
these horses for princely gifts, the other the short upright mane, and the
third the uniformly ash colour, so often recorded. If the first Duke of
Prussia, Albert von Hohenzollern, sent wild horses as highly esteemed gifts
to the mightiest sovereign of his days, the Emperor, and also to the Arch-
ORDER PERISSODACTYLA : ODD-TOED UNGULATES 333
duke Ferdinand and others, there can be no doubt that these horses were a
truly royal game like the Urus and Bison and not of a little valued feral
breed. And if the mouse-colour and the short mane are recorded for some
of the last survivors in Poland and Southern Russia, it must be almost
certain that there was at least a strong strain of true wild blood in these
horses. S. G. Gmelin, one of the many German explorers of Russia in the
days of the great Catherine, hunted these "Tarpans" in 1763 in the surround-
ings of Bobrowsk, Woronesh. After him the author named these horses
scientifically "Equus gmelini" but perhaps there is an earlier name: Equus
silvestris v. Brincken, dedicated to the mouse-dun wild horses of Poland,
surviving in the forest of Bialowieza until the middle of the eighteenth
century, and in another game park until 1812. Although protected very
strictly against poaching and illegal hunting, the wild horse in Prussia vanished
in the second half of the sixteenth century. In the well-known forest of
Bialowieza, Poland, the "Tarpanis" were hunted as royal game in 1409,
when King Wladislaw Jagiello arranged a great chase in honour of his cousin,
Witold of Lithuania. In the immense forests they survived until the eighteenth
century, when they were extirpated before the time of the famous hunting
of the Saxon Kings. Their last refuge in the Poland of to-day was accord-
ing to Vetulani the great game park of the Count Zamoyski, situated at
Zwierzyniec, near Bilgoraj. Here they were strictly protected, until in a
severe winter between 1810 and 1820, probably from 1812 to 1813, the feeding
was impossible. The last survivors were captured and given to the peasants
of the surrounding country. According to these facts there are in no other
district of Poland more typical "Tarpans" among the little horses of the
peasants than in the surroundings of Bilgoraj [c/. Janikowski, 1942]. Ve-
tulani has proved these Polish wild horses as a more or less degenerated
branch according to their being adapted to the unsuitable wood life of
the Eastern or Russian Tarpan.
The latter vanished from the fertile country of Woronesh before 1800, but
survived on the steppes of Tauria and Cherson until the middle of the nine-
teenth century. The last herds were certainly more or less intermingled with
feral horses, but the short mane being recorded even for the last example,
demonstrated the predominance of true wild blood. F. von Falz-Fein, the
well-known founder and owner of the matchless Ascania Nova Zoo, has
told the life-history of that last wild horse of Europe, an one-eyed old mare,
lingering for years around the feral horses of a certain Durilin, covered by
a domestic hehorse, captured, escaped with its filly, and some years later
hunted and killed on the ice by the peasants of Agaiman.
There is only one drawing from a living example hitherto known: in the
description of the travels of Gmelin, edited by Pallas after the tragic death
of his comrade. This picture, drawn by Borisow from a one year old mare,
was later on copied by Schreber in his "Naturgeschichte der Saugetiere."
It must be recorded that experiments for the rebreeding of the mouse-
dun Tarpan were started both in Germany and in Poland. In the Schorfheide
near Berlin and in the Munich Zoo the Germans try the rebreeding by
crossing the true yellow-dun Mongolian wild horse with mouse-dun mares
of various domestic breeds, while in Bialowieza the Poles settled upon some
most typical descendants of the last Bilgoraj wild horses, selected out of a
great number of peasant-horses in that district, without any interbreeding
of strange blood. The question is, which of the two trials will have the
better results.
The home of the mouse-dun Tarpan extended eastward over the river
Don and probably to the right bank of the Wolga. It is possible in earlier
334 EXTINCT AND VANISHING MAMMALS
days that these horses were also spread over the Caspian steppes, but as it is
difficult to distinguish the different records about other Equidae, e. g., the
Kulan, it is impossble to confirm that opinion.
The hillier steppe-country between the Wolga and the Ural-Mountains,
in the days of Pallas already crossed by a line of Kossak-posts, were roamed by
another wild horse. Pallas gives in his great travel-work, the well-drawn
portrait of a young filly, captured in the surroundings of Tozk then a little
Kossak post. That picture resembles in a high degree the Przevalski-fillies,
imported by Hagenbeck in 1899 and 1900. Together with the statements of
Pallas about the colour (Isabella to light bay), the "suberect" mane, the
tail, etc., there can be no doubt that these horses were almost as pure-bred
wild horses of the yellow-dun Przevalski-type as ever roamed the Dzungarian
Gobi. In the time of Pallas the wild horses were spread in scattered troops,
more or less intermingled with escaped domestic horses, over the steppes of
Western Siberia. Georgi, one of Pallas's fellow-workers, reports that they
were extirpated by a desolating horse-sickness in 1785 which destroyed also
the herds of the Kirghises and Kossaks, causing the death of about 85,000
horses. In 1876 the species was rediscovered by the great Russian explorer
Przevalski in the Dzungarian steppes south of Kobdo and named after him
by Poljakoff "Equus przevalskii."
Quagga
EQUUS QUAGGA Gmelin
Equus quagga Gmelin, Linnaeus' Syst. Nat., eel. 13, vol. 1, pt. 1, p. 213, 1788.
(South Africa.)
FIGS.: G. Edwards, Gleanings of Natural History, p. 29, pi. 223 (col.), 1758;
Ridgeway, Proc. Zool. Soc. London 1909, pp. 563-586, text-figs. 157-180,
reproductions of early figures and photographs of preserved specimens.
The vernacular name, Quagga, of this handsome zebralike species
is said to be derived from the Hottentot khoua khoua, in imitation
of its barklike cry. The Boers, however, often applied the same name
to the BurchelFs type of zebra, and it is sometimes loosely used for
that animal by writers.
Harris, whose folio Portraits of the Game and Wild Animals of
Southern Africa, 1840, provides some first-hand information on this
species, wrote that it stands 4.5 feet at the withers and has a total
length of 8.5 feet, but Cuvier (quoted by W. L. Sclater, 1900, vol. 1,
p. 295) gives the height at the shoulder as slightly less, about 4 feet
1 inch. The general ground color of head, neck, and body was dark
rufous brown or bay, becoming gradually more fulvous and fading
off to white behind and beneath. The midline of the back was
marked by a broad dark stripe. Against the background of bay, the
forehead was marked with longitudinal stripes and the cheeks with
narrow transverse stripes of buff, "forming linear triangular figures
between the eyes and the mouth." Muzzle black; neck and anterior
half of the body banded and brindled with creamy brown, broader
and more regular on the neck (extending across the short erect
mane), but becoming finally lost in spots and blotches on the rear
ORDER PERISSODACTYLA : ODD-TOED UNGULATES
335
half of the trunk. The legs, tail, and under surfaces were white,
with sometimes a short midventral dark line, and usually a black
spot behind the fetlocks. Individuals appear to have varied con-
siderably in the width and extent of the paler stripes and in the
amount of whitish on the rump, tail, and belly. These differences
have been made the basis of several subspecific names, but it is
now agreed that they are best considered as only variations of a single
species. Although Pocock earlier believed that the Quagga was
merely a southernmost form of the BurchelPs Zebra, and that certain
FIG. 34. Quagga (Equus qwagga). After Standard Natural History.
individuals nearly bridged the gap between the extremes, it has since
been shown that the two are doubtless separate species, and that the
Quagga, in addition to the well-marked color characters, was further
distinguished by cranial differences as well. For according to
Schwarz (1912b) the skull is the smallest of the three South African
species of zebras, and is characterized by its relatively wide zygo-
mata, narrower bony eye ring, broader forehead, greater separation
of the temporal ridges, the presence of a small suborbital pit, and
by having the posterior border of the nasals heart-shaped. While
such characters may be subject to individual variation, they may for
the present be regarded as valid.
The Quagga seems first to have been brought to the notice of natu-
ralists by George Edwards, who in his Gleanings (1758) published
336 EXTINCT AND VANISHING MAMMALS
a colored plate of a female, which he supposed to be the female of the
Mountain Zebra. Buffon in 1782 (Hist. Nat., Suppl, vol. 6, p. 85)
was the first to give an authentic account of the animal on the basis
of notes supplied him by Allamand, from Colonel Gordon, a South
African resident. Subsequently Edwards' description became the
basis of Gmelin's name Equus quagga. So far as the records show,
the Quagga always had a somewhat restricted range and was con-
fined chiefly to the southeastern corner of the Cape region, from
Algoa Bay westward at least to Prince Albert (where Barrow reports
it in 1801), and Swellendam, some 100 miles east of Cape Town,
where Sparrman first saw it, northward to the Orange Free State and
the Vaal River, and coastwise to the Kei River. There is practically
no record of travel in South Africa between the days of Kolben in
1705 and the visit of the Swedish naturalist Sparrman in 1775. The
latter, however, found the Quagga as near the coast of Algoa Bay as
Uitenhage. He secured a foal which is still preserved in the Riksmu-
seum at Stockholm. Quaggas were apparently still plentiful in the
first quarter of the last century, especially in the districts of Aberdeen
(Lichtenstein, in 1804) and in Fraserburg and Hanover where Bur-
chell in 1812 found them abundant, in troops of 30 to 50 on the plains.
These made an impressive sight traveling in single file as was their
curious habit, or when startled, wheeling in unison like a squad of
cavalry. It was said that they frequently associated on the plains
with the White-tailed Gnu or with ostriches; whereas the Burchell's
Zebras preferred the companionship of the Brindled Gnu. Where,
as in the Orange Free State, the range of the Quagga met or over-
lapped that of Burchell's Zebra, it is said the two did not mingle.
The Boer farmers evidently took heavy toll of them in these years,
greatly reducing their numbers, and using the meat to feed their
workers although themselves preferring more tasty kinds of game.
The result of this constant persecution was that by the late 50's the
Quagga was practically extinct south of the Orange River. According
to Bryden (1889) the last known instance was of two shot in 1858
near the Tygerberg, a solitary mountain rising abruptly from the
plains near Aberdeen. His informant, the successful hunter, recalled
the affair well. Farther north, however, in the Orange Free State,
Quaggas were still numerous. About 1865, the Boers of this state
began the exploitation of the large game of the region for their hides.
With characteristic industry and deadly skill they gathered and
shipped to the coast hides by the wagonload, among which those of
Quagga and Burchell's Zebra were especially in demand. They also
made use of Quagga hides for grain sacks, and Bryden (1889) men-
tions seeing old Quagga-skin sacks still in use at the time of his visit.
The exact date of the final extermination of the Quagga is unknown,
but it is generally believed that the species continued well into the
ORDER PERISSODACTYLA I ODD-TOED UNGULATES 337
'seventies in the Orange Free State, probably, according to W. L.
Sclater (1900) , "till 1878 at least," but he adds, "it is difficult to
obtain any accurate information on the subject, as in so many cases
this and BurchelPs Zebra are confused together, especially as they
were both known under the name of quagga."
In the earlier days of the last century and even shortly before,
Quaggas were occasionally tamed and also exported alive to the
zoological gardens of Europe. In disposition it was said to be much
more tractable than the BurchelPs Zebra, in captivity quickly be-
coming docile and tamable. On various occasions they were broken
to harness, and Sir William Jardine even mentions that a Mr. Sheriff
Parkins early in the nineteenth century drove a pair in London,
and was often seen in Hyde Park riding in a phaeton after them.
Probably one of the first Quaggas to reach Europe alive was the one
belonging to His Royal Highness the Prince of Wales, from which
in 1751 George Edwards made his colored drawing. The specimen
now in the Paris Museum was brought to the menagerie of the King,
at Versailles, in 1793. Others were later imported by animal dealers
such as Frank at Amsterdam. Of the various specimens extant in the
museums of Europe, the larger part were brought in alive and
received by the museums after having died in captivity. Thus the
locality of capture is in most of these cases unrecorded. In 1858, Sir
George Grey presented to the Zoological Society of London a male
Quagga which died six years later, in 1864. "It is the mounted skin,
skull and skeleton of this male which is now in the British Museum"
(Ridgeway, 1909). Previously in 1851 the Society had purchased
a female Quagga which survived in Regent's Park, until 1872, ap-
parently nearly the last living example of the species of which any
positive record exists. Further, this was the only living Quagga ever
to be photographed, and the picture has been reproduced by Ly-
dekker in his Guide to the Specimens of the Horse Family and by
Ridgeway in his paper of 1909. The skin was not in condition to be
preserved; but it is said that the skeleton was saved and mounted,
although at the present time it has been lost sight of and is evidently
not the one now in the British Museum. Finally, the last known
living specimen seems to have been one that died in the Berlin
Zoological Garden in 1875. The skin is mounted in the Zoological
Museum in that city, and the skeleton is also preserved there.
Combining the lists of Ridgeway (1909) and Hilzheimer (1912),
the known specimens of the Quagga in the museums of the Old World
are the following (arranged alphabetically by location) :
1. Amsterdam Museum. Mounted specimen, and separate skull. Figured
by Lydekker (Proc. Zool. Soc. London, 1904, vol. 1, p. 430, text-fig. 86) and
by Ridgeway (1909, p. 579, text-fig. 170).
2. Basle Natural History Museum. A mounted female from Silo (Shiloh),
12
338 EXTINCT AND VANISHING MAMMALS
Cape Colony, presented in 1864. Figured by Ridgeway (1909, p. 565, text-
fig. 157).
3. Berlin, Zoological Museum. Mounted female, that died in the Berlin
Zoological Gardens in 1875; also its skeleton and two other skulls. Figured by
Ridgeway (1909, p. 578, text-fig. 168).
4. Cape Town Museum, South Africa. Mounted foal, from Beaufort West,
about 1860. Figured by Ridgeway (1909, 580, text-fig. 171).
5. Darmstadt Museum. Mounted specimen.
6. Edinburgh, Royal Scottish Museum. A mounted specimen, purchased by
the University of Edinburgh during the year ending June 1818. Figured
by Ridgeway (1909, p. 575, text-fig. 165).
7. Elgin Museum, Scotland. A mounted head and neck, from King Wil-
liam's Town, 1861. Figured by Ridgeway (1909, p. 581, text-fig. 172).
8. Frankfurt a.M., Senckenberg Museum. A well-mounted skin and its
skull, received in 1831 by exchange with the Leiden Museum.
9. Leiden, Dutch State Museum of Natural History. Mounted male and
its skeleton, shot near Steenbergen, June 15, 1827. Figured by Ridgeway
(1909, p. 577, text-fig. 166).
10. London, British Museum. Ridgeway (1909, p. 574) has cleared away
the confusion regarding the number and origin of the specimens in this
Museum. Apparently the only one is a male skin, mounted, and the skeleton
of the same animal, which had been presented to the Zoological Society of
London by Sir George Grey in 1858 and lived in the Society's Gardens until
its death in 1864. The female Quagga which lived in the Gardens from 1851
to 1872, was photographed in life, but its skin, upon its death, was in too poor a
state to be preserved. Its skeleton, however, was mounted but cannot now
be traced. The male specimen as mounted is figured by Ridgeway (1909,
p. 573, text-fig. 163) and the living female is figured by Ridgeway (1909, p. 575,
text-fig. 164) from York's photograph of the animal.
11. Mainz Museum, Germany. According to Hilzheimer (1912) there were
four mounted Quaggas in this collection, but Schwarz (1912) who also examined
them, asserts that one of the four is a Burchell's Zebra.
12. Munich Natural History Museum. A mounted specimen purchased in
1835, and a separate skull that may or may not belong to the same individual.
It was this specimen that was the original of the figure by Wagner in "Schreber's
Saugthiere, Supplement." Figured by Ridgeway (1909, p. 579, text-fig. 169).
13. Paris Museum of Natural History. A mounted specimen. According to
the communication of Dr. E. L. Trouessart, it was received living after the
institution of the Museum's menagerie from the old menagerie of the King,
at Versailles in 1793. No more precise locality is given for it than "Cape of
Good Hope." Figured by Ridgeway (1909, p. 577, text-fig. 167).
14. Stockholm, Riksmuseum. The mounted specimen here is a full-grown
fetus, brought back by the Swedish traveller Sparrman in 1775. It is therefore
the oldest extant specimen and appears to have the pale stripes clearer and
extending farther back than usual. While the exact locality is unrecorded,
Sparrman mentions that he first saw Quaggas at Swellendam. Figured by
Ridgeway (1909, pp. 570, 571, text-figs. 160, 161), both from a photograph and
from a recent painting.
15. Stuttgart Museum, Germany. According to Hilzheimer (1912) this
museum contains a skull and footbones of the Quagga. He further mentions
that in the Stuttgart Altertums Collection is a miniature model of a Quagga,
of which he gives a figure, and suggests that it was probably prepared from
the two animals which Frederick I had in his menagerie in 1812-16. It may
therefore have a certain authenticity.
ORDER PERISSODACTYLA : ODD-TOED UNGULATES 339
16. Tring Museum, England. A mounted specimen, interesting for the
distinctness of the posterior stripes. It was described and figured by P. L.
Sclater (Proc. Zool. Soc. London, 1901, vol. 1, p. 166) who believed that it
was the same as the animal formerly living in the London Zoological Gardens
from 1851 to 1872. That this, however, is not the case was made clear by
Ridge way, who publishes a letter from E. Gerrard (who sold the specimen
to Lord Rothschild) stating that he had purchased the animal as an old
mounted specimen from a Mr. Frank of Amsterdam and had remounted it
before selling it to the Tring Museum. Figured by Ridgeway (1909, p. 569,
text-fig. 159).
17. Turin, Zoological Museum. A mounted female, and its skull. The
specimen was purchased in 1827 from the English dealers, Leadbeater father
and son. It was made the type of Equus trouessarti, figured and described
by Camerano (1908, pi.).
18. Vienna Museum. A mounted female, procured by Ecklon, in 1836.
The specimen was described by Lorenz (Proc. Zool. Soc. London, 1902, vol. 1,
p. 32) and figured from a photograph which is again reproduced by Ridgeway
(1909, p. 568, text-fig. 158).
19. Wiesbaden Museum, Germany. A mounted specimen, male, which was
bought in 1865 from Frank, the Amsterdam dealer. It has no more definite
locality than "South Africa." Figured by Ridgeway (1909, p. 572, text-fig. 162).
From this enumeration it appears that there are in the museums of
Europe 17 mounted skins (one a fetus) , a mounted head, 3 skeletons,
and 7 skulls ; while elsewhere the only known specimen is a mounted
skin of a foal, in the South African Museum.
G. M. A.
Burchell's Zebra or Bontequagga
EQUUS BURCHELLII BURCHELLii (Gray)
Asinus burchellii Gray, Zool. Journ., vol. 1, p. 247, 1824. (Little Klibbolikhoni
Fontein, Bechuanaland, South Africa.)
FIGS.: Gray, op. tit., pi. 9, figs. 1, 2; Lyon, M. W., Jr., Proc. U. S. Nat. Mus.,
vol. 32, pis. 1-3, 1907 (photographs of mounted specimen) ; Cabrera, Jour.
Mamm., vol. 17, p. 97, figs. 1-5 (pattern diagrams), 1936; Pocock, Proc.
Zool. Soc. London, vol. 1, p. 485, fig. 48, 1909 (photo.).
Although, on account of its somewhat variable pattern, the Bur-
chell's Zebra has been divided into many nominal races, only four of
these are regarded as valid by Cabrera (1936) in his recent review.
Of these four, the typical burchellii is now extinct, and there are few
specimens preserved.
About the size of a small horse, with erect mane and tufted tail,
the color pattern consists of alternating dark-brown or black stripes
and whitish stripes. Of the former, there are about ten on the neck,
a vertical shoulder stripe, four body stripes, then on the flanks three
or four that turn back dorsally, somewhat paralleling the median
black stripe. The characteristic feature of typical Burchell's Zebra
is that the lower haunches and both fore and hind legs lack the small
340
EXTINCT AND VANISHING MAMMALS
transverse stripes that are increasingly developed in the more
northern races, and the tail is white.
The former range of this race was rather limited and covered what
is now the Orange Free State and southern Bechuanaland, in South
Africa, but apparently did not extend to the south of the Orange
River. Over the plains of this region it once abounded "in countless
thousands," but with the coming of white hunters, followed by
settlers in the Orange River colony, it had already become rare by
the middle of last century. Many were exported to Europe for
FIG. 35. Burchell's Zebra (Equus burchellii burchellii). After Brehm.
zoological gardens, and it is in part from these that have come the
few specimens still preserved in museums. There is a specimen in the
British Museum, one in the Tring Museum, and a third in the
Bristol Museum, in England, and there is a mounted one in the U. S.
National Museum, and one in the Paris Museum, with a few others in
other museums, as Berlin, Leiden, and South Africa. The last
living specimen, so far as known, was one kept in the London
Zoological Gardens, where it was received apparently in 1909, after
evidently having been in captivity for a period.
From Benguela west to Southern Rhodesia and Zululand, this race
is replaced by the race antiquorum, with more cross-striping on the
upper parts of the limbs. At the present time this animal still exists
in some numbers in the west of South-West Africa. From the Lim-
popo River northeastward to the Loangwa and Rovuma Rivers is
ORDER PERISSODACTYLA : ODD-TOED UNGULATES 341
found the race selousii, distinguished by "having the limbs striped to
the coronet, and the body with numerous narrow stripes and few and
faint shadow stripes." Still farther northeastward, is the race bohmi,
inhabiting the plains country of eastern Africa north to Lake Rudolf,
in which the shadow stripes (between the clear black stripes) are
absent and the haunch stripes broad and black, and the limbs cross-
striped nearly to the hoof. This race is still common.
The chief enemy of the zebras, apart from man, is the Lion, which
seems specially fond of zebra meat and finds it easily obtainable.
The zebras go in herds which may at times be of large size. Fre-
quently Gnus, of one species or another, associate with these herds,
as if for companionship. Zebras have at times been tamed and
trained to harness but are of rather uncertain disposition and no
great use of them in this way has been made. Their meat, though
relished by the natives, is not popular among white hunters, accord-
ing to Selous. The hide is often used as leather. In regions where
agriculture is practiced, Zebras often become a nuisance to the
ranchers by stampeding and breaking through barbed-wire fencing.
They are said to have a good deal of curiosity, and especially where
mules or donkeys are in camp, will often approach closely to survey
them. According to Major Flower, they often in captivity live over
12 years, and have been known to reach 28 or 29 years in zoological
gardens. The name "quagga" is said to be derived from the noise
they make, a sort of honking bark. They are much dependent on
water, and drink at least once a day, often at night, but approach
the waterhole with caution, for fear of lurking lions.
Although, with the exception of the typical burchellii, none of the
races is in present danger of extinction, their numbers will undoubt-
edly diminish except in areas where large extents of grasslands
as in East Africa afford them range.
G. M. A.
Mountain Zebra. Wildepaard (Boer). Dauw (Hottentot)
EQUUS ZEBRA ZEBRA Linnaeus
Equus zebra Linnaeus, Systema Naturae, ed. 10, vol. 1, p. 74, 1758. (Probably
the Drakensberg and other mountain ranges, Cape of Good Hope.)
Hartmann's Mountain Zebra
EQUUS ZEBRA HARTMANNAE Matschie
Equus hartmannae Matschie, Sitzber. Ges. Naturf. Freunde Berlin, p. 174,
1898. (Between Hoanib and Unilab Rivers, South-West Africa.)
FIGS.: Of typical form: Lydekker, 1912, pi. 20, fig. 1; J. E. Gray, Knowsley
Menagerie, pi. 56, 1850; Pycraft, 1936, p. 850, fig. Of E. z. hartmannae:
Haagner, 1920, fig. 66; Zukowsky, 1924, fig. 6; Maydon, 1932, pi. 125;
Shortridge, 1934, vol. 1, pi. opp. p. 389; Pocock, 1937, p. 717, fig.
342 EXTINCT AND VANISHING MAMMALS
Since there is still some doubt (Shortridge, 1934, vol. 1, p. 389)
whether the Mountain Zebra of the dry South-West African uplands
is recognizably distinct from the typical E. zebra, the two may be
treated together, pending further study of specimens. Also some
authors, among them Captain Shortridge (op. cit.), prefer to regard
this as a genus distinct from Equus or from the South African
Quagga, calling it Hippotigris. Since in skeletal and tooth characters
it is very little different from the horses, a conservative course is to
regard the latter as a subgenus of Equus.
The Mountain Zebra was the first of the zebras known to
Europeans and is the smallest of the three species, standing about
12 hands high (48 inches) at the shoulder. Length of head and body
7 feet 4 inches; tail, with terminal hairs, 23 inches (W. L. Sclater,
1900). Sclater gives the following description:
Body, head, and limbs closely covered with black or almost black stripes,
broader than their white interspaces; on the face the dark markings below
the eyes become reddish passing into large nostril patches of the same color,
but the muzzle itself is black; . . . ears long and rather narrow, posteriorly
the basal two-thirds striped, the terminal third black, the extreme point
white; . . . longitudinal dorsal stripe only noticeable over the haunches,
transverse stripes of the barrel extending back over the haunches to the
base of the tail forming here the so-called gridiron pattern; no shadow
stripes; hairs along the back to the shoulders reversed; belly white, except
for a longitudinal dark band running along its anterior portion which is
never reached by the transverse body stripes; limbs transversely marked
down to the hoofs, . . . the pasterns being quite black; . . . hoof rather
narrow, compact and solid; tail reaching the hocks with a median black line
and traces of transverse bars at the base; the distal quarter with a tuft of
long black hairs.
Hartmann's Zebra is believed to differ from the typical race in
its larger size and more widely spaced stripes, so that the pale stripes
are equal to or even slightly wider than the black ones. The legs
"are almost evenly banded black and buff the black not predomi-
nating as in zebra." However, this pattern varies individually and as
yet it is uncertain whether the characters claimed are relatively
constant.
As its name implies, this zebra was an upland species, living in
the mountains, "from Great Namaqualand (and possibly Damara-
land), through the various ranges of Cape Colony to the Great
Drakensberg chain, and thence to the end of that range. ... At
the present day," wrote Bryden (1899, p. 94) , "it is only to be found
in small troops here and there in Cape Colony. It is very doubtful
whether any now remain in Great Namaqualand, where, sixty years
ago, Sir James Alexander found them in considerable numbers. It is
probable that the Hottentots . . . , who are excellent shots . . . ,
have destroyed the last remnants ... in ... Great Namaqualand.
ORDER PERISSODACTYLA I ODD-TOED UNGULATES
343
In Cape Colony, where these zebras are, as far as possible, preserved,
small troops are to be found in the mountains of the Sneeuwberg,
Witteberg, Tandtjesberg, Zwartberg, the Winterhoek, and one or two
other ranges. A few still linger along the Drakensberg. . . . Near
Cradock, . . . only a few years since, a troop of twenty was seen."
Under date of January 23, 1935, Herbert Lang, the well-known
explorer and authority on large game mammals of South Africa,
FIG. 36. Mountain Zebra (Equus zebra zebra). After photographs in
Brehm and Newnes.
wrote that "there must be still about a hundred Mountain Zebras
in various places in spite of the reports to the contrary. In 1926
I traveled through all these regions to observe these zebras in their
haunts. One must have seen these herds of Mountain Zebras on
their actual trails to be enthusiastic about them and their protec-
tion. There can hardly be anything more fascinating." By 1937,
according to editorial notes in the Journal of the Society for the
Preservation of the Fauna of the Empire, a census of these zebras
showed the following 45: in the Oudtshoorn Area, near George, 20
on the farm of Peter. Heyns ; in the Cradock Area, 10 on the farm of
Paul Michau, 8 on the farm of Lombard, 7 on that of Osborn. The
344 EXTINCT AND VANISHING MAMMALS
proportion of sexes among these animals was believed to be very
uneven with "a great shortage of mares." This number is still less
than a few years previously, for E. L. Gill, writing in December 1932,
said that at Cradock there were "two or three small herds which seem
likely to die out. The Oudtshoorn herd on the farm Mount Hope,
has been carefully preserved by the owners, the brothers Heyns, and
is still flourishing. It numbers somewhere about 70 animals and con-
stitutes the chief hope for the survival of the species."
Efforts have been made at various times in recent years to induce
the Government to purchase a portion of the Mount Hope Farm as a
permanent Mountain Zebra reserve. In 1937, the Government at
last voted to ask the Parliament of the Union of South Africa for
7,600 for the establishment of such a reserve and hopes are high that
it may actually be created, before it is too late. Notwithstanding
that the species is protected at all times by the South African
Government, the actual enforcement of this protection has in the
past been difficult. It is one of the species listed for complete pro-
tection by the London Convention of 1933.
While "the advance of civilization" is blamed for the reduction
in numbers of this zebra, no doubt much blame must also be laid
upon the native and white poachers with modern rifles, as well as
to other methods of extermination. Bryden (1899) wrote that occa-
sionally the weather is so "severe among the Cape mountains that
even the tough zebra succumbs" and that in "the old days in Cape
Colony, the Boers were in the habit of hunting these animals for the
sake of their hides and of capturing the young alive for the purpose
of being broken to harness." For in the last century "a fashion for
using Mountain Zebras in harness seems ... to have sprung up in
the Mauritius, and ... a good many of these animals were exported
from the Cape to meet the requirements of the French colonists.
... A premium of 20 was at the same time offered for the young
of these animals delivered in Cape Town." "The Boers, to save
themselves the trouble of shooting, occasionally succeeded in driving
a number of these animals over the edge of a precipice, thus securing
the skins at their leisure" (Bryden, 1899) .
Although Bryden (1899) feared that within the "next fifty years
this zebra will have joined the ranks of extinct creatures," there
seems still some hope of preserving a remnant, owing chiefly to the
interest of those farm owners on whose lands the survivors still hold
out, and an awakened enthusiasm on the part of the Government to
do what it can.
While the future of the Mountain Zebra in the Cape Province is
none too rosy, it still occurs in the form hartmannae in small numbers
among the mountain ranges of the western and northwestern parts
of South-West Africa and northward across the Cunene into south-
ORDEK PERISSODACTYLA I ODD-TOED UNGULATES 345
western Angola at least as far as Elephant Bay, 100 miles north of
Mossamedes, where, however, it is not found more than 30 miles
inland. Shortridge (1934, vol. 1, pp. 390-396) has gathered together
the available information concerning it in this region. The eastern
limits are found in the Kaokoveld about a hundred miles from the
west coast. Here it is sometimes found in association with the Bonte-
quagga (E. burchellii antiquorum) but is much fewer in numbers. It
is partial to the crests of arid gorges, and its small cupped hoofs
are adapted for rough country. Large numbers are said to be found
in the Omaruru and the Maltahohe districts, but elsewhere they are
less common. Steinhardt saw them digging for water in sandy river
beds of this arid country, making pits half a meter deep. They may
not drink regularly but sometimes keep away from water as long as
three days. They are shy and suspicious and difficult to approach
under usual conditions. The Cape Mountain Zebra is believed to be
a slow breeder, with foal every second year or so. The period of
gestation is said to be about twelve months. The height at the
shoulder is said to be in hartmannae about 52 to 54.5 inches, hence
somewhat taller than the typical race.
G.M. A.
Nubian Wild Ass. Nubischer Wildesel (Ger.)
ASINUS ASINUS AFRICANUS Fitzinger
Asinus ajricanus Fitzinger, Naturgesch. Saugethiere, vol. 3, p. 667, 1857.
(Lydekker (1916, vol. 5, p. 38) gives the type locality as "Nubia (accord-
ing to Matschie, Erythraea) .")
SYNONYM: ? 'Asinus asinus dianae Dollman (1935).
FIGS.: Proc. Zool. Soc. London 1884, pi. 50, fig. 2, 1885; Lydekker, 1904,
pi. 20, 1912, pi. 20, fig. 2, and 1916, vol. 5, p. 37, fig. 16; Antonius, 1929,
p. 290, fig. 1; Zammarano, 1930, p. 87, fig.
This subspecies is "by no means common" (Brocklehurst, 1931,
p. 15).
"General colour of upper-parts greyish-fawn, with the muzzle, a
broad ring round each eye, . . . and the under-parts, white or
whitish ; the legs being of the same pale hue, with some greyish on the
front surface, and a few small dark spots on each side of the fetlocks.
The mane ... is short, upright, and dark brown or blackish ....
The narrow dorsal stripe ... is continued as a thin line well on to
the tail"; the two branches of the shoulder stripe are about 5-6
inches in length. "The long hairs of the terminal tail-tuft ... are
mingled black and grey. The ears are about 10J in. in length, and
are black at their tips .... On the inner side of the lower part of
the fore-leg is a chestnut patch." Height of male at shoulder, 45^ to
47i inches. (Lydekker, 1904, p. 594.) Baker (1867, p. 56) gives the
height of a male from the Atbara River as 55-56 inches.
346
EXTINCT AND VANISHING MAMMALS
A. a. dianae Dollman (1935), from south of Tokar near the
Eritrean boundary, is so weakly differentiated from ajricanus as to
seem scarcely worthy of recognition ; it may represent a slight inter-
gradation toward somaliensis.
"The Nubian wild ass ... inhabits , . . Sennar and Nubia, its
range formerly extending as far as the fifth cataract of the Nile
FIG. 37. Nubian Wild Ass (Asinus asinus ajricanus)
.... Year by year the range of this race appears to become more
restricted; and unless protective measures be taken, there is danger
that it may be exterminated." (Lydekker, 1908, p. 66.)
Heuglin (1861, p. 19) reports Wild Asses as occurring from Suakin
to the Nile at Berber, in all northeastern Sennaar, and in the plains
of the Barka River. He says he met with them commonly about
the ruins of Wadi Safra, then on the Atbara, and along the route
from Taka toward Suakin ; and during the rainy season they appear
as far north as the Desert of Korosko. (Korosko is on the Egyptian
part of the Nile, at about lat. 22 30' N.; but Flower remarks (1932,
p. 432) : "There appear to be no certain records of genuine wild
asses having occurred in Egypt during the nineteenth century.")
ORDER PERISSODACTYLA ! ODD-TOED UNGULATES 347
Sir Samuel Baker (1867, pp. 55-56) writes concerning the Wild
Ass along the Atbara River: "Those who have seen donkeys in
their civilized state have no conception of the beauty of the wild
and original animal. . . . The animal in its native desert is the per-
fection of activity and courage; there is a high-bred tone in the
deportment, a high-actioned step when it trots freely over the rocks
and sand, with the speed of a horse when it gallops over the bound-
less desert. No animal is more difficult of approach; and, although
they are frequently captured by the Arabs, those taken are invari-
ably the foals, which are ridden down by fast dromedaries, while
the mothers escape."
Matschie gives (1894, p. 73), as the northernmost locality, the
Wadi el Homar, a little north of Berber.
"Their flesh is eaten by the Arabs of the Soudan. They are
ordinarily met with in twos and threes, or small herds." (Bryden,
1899, p. 70.)
"This animal is found at the foot of the Gebel Hennah, near
Tokar. It is common in the Khor Sabbat parallel to the Khor
Baraka. Captain O'Connor informs me that he has often seen them
at the Khor Sabbat, on the plain of Tokar." (Anderson, in Ander-
son and de Winton, 1902, p. 330.)
"Neither the wild asses nor the zebras of Africa are pursued with
much enthusiasm by sportsmen, and the first-named animals are
so shy and wild that whilst it is very difficult to get within shot of
them on foot, if they are hunted on horseback they are so fleet and
enduring that they can only be overtaken with great difficulty even
by a really fast horse. . . . Thus the wild ass is seldom shot, and
is probably of less interest to the average sportsman than any other
African game animal." (Selous, 1914, p. 36.)
"The Wild Ass is found in the Sudan in the neighbourhood of the
Atbara River in the provinces of Berber and Kassala; it is also
found in the Red Sea Province south of Suakin.
"They have been strictly protected for a number of years, and
although by no means common there is not, at present, any danger
of their being exterminated." (Brocklehurst, 1931, p. 15.)
Some years before 1932 Wild Asses were fairly common in the
region of the Baraka Wadi near the border of Eritrea (Maydon,
1932, p. 203) .
In all probability the very few Eritrean specimens now extant live
in the region- of Upper Barca (De Beaux, 1935, p. 12) .
Powell-Cotton (in Dollman, 1935, p. 134) writes concerning Wild
Asses south of Tokar near the Eritrean boundary: "Beween 18 Feb-
ruary and 2 March 1934 we saw the animals on four occasions, as
follows, 1, 2, 2, 3, and secured the two specimens permitted us.
. . . The Arabs . . . leave the Wild Ass unmolested as they do not
348 EXTINCT AND VANISHING MAMMALS
eat its flesh, and they told us they were more numerous across the
Eritrean boundary."
Antonius (1938, p. 560) writes:
[The Nubian Wild Ass] until recent times spread over the mountainous
semi-deserts of Nubia and the Eastern Sudan from the Nile to the shores
of the Red sea. The last specimens of Nubian origin known to European
observers were two males shot by Sir Reginald Loder about 1925, near the
Gebel Raboba, on the Erythrean frontier, and an old female, living in the
Zoological Gardens at Rome for many years, the photograph of which is
reproduced by Zammerano in his "Fauna e Caggia." To-day there is a living
specimen in the New York Zoological Park, and also a stallion in the Zoo
at Rome. The Nubian ass was domesticated by the ancient Egyptians at
least in 3500 B. C., and being hunted by the King Rameses III. as is shown
at the temple of Medinet Habu it was well known as a wild animal in
Egypt until at least 1100 B. C. . : .
The so-called "Nubian Asses" shown in many Zoos, are by no means pure
bred descendants of the true wild stock; therefore it seems nearly certain
that the Nubian ass is to-day as thoroughly extinct as his Atlantic cousin.
[This remark on extinction is somewhat premature, in view of the statements
just quoted from Brocklehurst, Maydon, De Beaux, and Powell-Cotton.]
All forms of the Wild Ass in Africa enjoy complete protection
under the London Convention of 1933.
Domestication. According to Lydekker (1916, vol. 5, p. 37), the
typical subspecies of the African Wild Ass (A. a. asinus) is repre-
sented by the Domestic Ass of Asia. He also states (1912, p. 217)
that the wild animal was first tamed in the Mediterranean countries.
This ancestral wild animal was doubtless distinguishable from the
present domesticated animal, but whether it was identical with any
of the wild forms now recognized (africanus, atlanticus, or somali-
ensis), it is impossible to say. Werth (1930, p. 351) suggests Abys-
sinia as the place of first domestication. If, however, the wild ances-
tor inhabited the eastern Mediterranean region, where no wild repre-
sentative of Asinus asinus now exists, it may have differed from any
of the currently recognized forms. In the account of Asinus hemionus
hemippus (p. 368) I have suggested the possible occurrence of some
wild form of Asinus asinus in the Palestine region within compara-
tively recent times.
While space does not suffice to discuss the subject of domestication
at any length, the following quotation from Peake (1933, pp. 98-99)
is apropos here:
The ass was used at a very early date, both in Egypt and in Mesopotamia,
but it is impossible to say at present to which region to ascribe the priority.
Towards the close of the predynastic period in Egypt, just before 3400 B. C.,
the Libyan tribes, who dwelt in the desert to the west of the Nile Delta,
possessed large herds of asses, and this indicates that this animal had been
known to and possessed by them for a long time. At Ur, in Mesopotamia,
asses were used by those kings and queens who were buried in the famous
death-pits, accompanied by their slain retainers, and these must be relegated
ORDER PERISSODACTYLA I ODD-TOED UNGULATES 349
to quite as early a date, and are probably much earlier. All our evidence
goes to show that asses had been tamed, and were used as beasts of burden,
both in Egypt and Mesopotamia, between 4000 and 3000 B. C., and may have
been domesticated at a considerably earlier date.
On the island of Sokotra there are herds of wild or feral asses
closely resembling A. a. africanus in color but of smaller size (stand-
ing from 38 to 40 inches at the shoulder) . They are regarded as "the
survivors of Nubian ancestors brought from the Red Sea coast by,
probably, the ancient Egyptian incense collectors." Their introduc-
tion is presumed to date back "for some thousands of years" per-
haps a sufficient length of time for their insular habitat to have pro-
duced degeneration in size. "As they are never shot at and rarely
molested by the natives, they were by no means wild." (C/. H. 0.
Forbes, 1903, pp. xxxviii, 6, 9-11, pi. 2.)
[There has been much discussion as to the possible existence of
Wild Asses in the Sahara. Tristram (1860, p. 318, as quoted by
Blyth, 1862, pp. 363-364) "heard that wild Asses were to be occa-
sionally found in the Soufa desert, on the route to Ghadames." He
was shown one that "had been caught when very young, and was
considered unusually tame for one of his species." It was "of a rich
slatish ash-colour"; dorsal stripe and shoulder stripe present; nose
and limbs white; mane and tail blackish. The adults were said to
be "very difficult to entrap and impossible to train."
On a journey through the Tuareg country in 1913-14 Geyr von
Schweppenburg (1917, p. 298) learned that wild or feral asses were
not uncommon there in previous years, but by that time had virtually
disappeared, having been captured, shot, and fed to dogs. The local
Mohammedans did not eat the flesh. He was inclined to consider
that at least some of the asses were genuinely wild, and not merely
feral. He also mentioned reports of Wild Asses in this region by
previous travelers (Duveyrier, Bissuel, Benhazera, and others).
According to Spatz (in Werth, 1930, p. 347) , these wild or feral
asses constantly molested the domesticated animals and often led
them astray.
Antonius (1931, pp. 133-136, pi. 3, fig. 3) calls attention to some
wild-living asses reported by Fraulein von Wagner-Jauregg in the
Hoggar massif in the southern territories of Algeria, where the
Tuaregs distinguish them by name from the Domestic Asses. A
captured foal showed evidence of a strong wild-blooded component,
even if it coud not be considered a pure-blooded wild ass. It is feared
that these wild-living asses of the Hoggar will disappear if energetic
protective measures are not adopted.
On the other hand, Hilzheimer and Spatz (Zeitschr. fiir Sauge-
tierk., vol. 17, p. 15, 1932) express the opinion that the animals of
the Hoggar are merely Domestic Asses that have run wild. Selous
350 EXTINCT AND VANISHING MAMMALS
(1914, p. 35) and Lavauden (1933, p. 20) state that Wild Asses do
not pass west of the Nile.
Antonius (1938, p. 560) says:
It is very interesting that the only mention of a wild ass captured in the
Western Sahara, given by Canon Tristram, agrees not with the appearance
of the Atlantic, but with that of the Nubian, race. Although very exactly
describing the coloration of his wild born ass, Tristram does not say any-
thing about banded limbs. The wild asses once roaming in the neighbour-
hood of the caravan route to Chadames [Ghadames] and Chat [Ghat] were
therefore probably not of the strongly marked Atlanticus-type, but of the
Nubian with unbanded limbs. This opinion is strengthened by the colours
of the so-called "ahoulil" of the Tuareg wild or feral asses, lingering in the
Ahaggar mountains in single pairs and small troops. A filly of these ahoulil,
captured in 1927 in the heart of the Ahaggar mountains for the Schonbrunn
Zoo, but dying before transported to Europe, was of exactly the same Nubian
type. Probably there is in these ahoulil at least a strong strain of originally
wild blood, more or less intermingled with the Atlanticus blood of escaped
domestic Tuareg donkeys.
Finally comes the extremely interesting information from Mai-
brant (1936, p. 27) that in French Central Africa Wild Asses are
restricted to the massif of Tibesti. Views differ as to their origin and
systematic status, but Malbrant inclines toward the opinion that
they are genuinely wild animals of the subspecies africanus. They
exist in the region of Zouar and, farther north, in the Tarsoa, moun-
tainous ridges situated north of Emi Koussi. Here they are not rare.
The plateau of Daski and the region of Trotron (between Yebi and
Zoumri) likewise shelter many. They live in bands of as many as
30 or 40 individuals. The natives capture young ones in snares near
the water-holes, train them, and use them as pack animals.
A view differing from Malbrant's is held by Thesiger, who remarks
(1939, p. 441) that in Tibesti "donkeys are extensively used, and
many have run wild among the mountains probably for generations."
The solution of the problem of the Tibesti Asses awaits the collec-
tion of specimens.]
Somali Wild Ass. Somali- Wildesel (Ger.)
ASINUS ASINUS SOMALIENSIS Noack
A[sinus] taeniopus var. Somaliensis Noack, Zool. Garten, vol. 25, no. 4, p. 101,
1884. ("Somaliland"; type locality restricted by Lydekker (1916, vol. 5,
p. 39) to "Berbera district of [British] Somaliland.")
FIGS.: Nouv. Arch. Mus. Hist. Nat. Paris, vol. 5, Bull., pi. 5, 1869 (subsp.?) ;
Proc. Zool. Soc. London 1884, pi. 50, fig. 1; Akeley, 1914, pp. 112, 115, 117,
figs.; Zammarano, 1930, p. 88, fig.; Schmidt, 1938, pi. 11.
In British Somaliland this Wild Ass "is strictly preserved, but . . .
much reduced in number" (Antonius, 1938, p. 560) .
It is more strongly built than A. a. africanus] general color a deli-
ORDER PERISSODACTYLA .' ODD-TOED UNGULATES 351
cate reddish ash-gray ; snout gray ; behind it a broad, light gray band
from the nose to beyond the corner of the mouth ; a light ring about
the eye ; inner surface of ears ash-gray, with black border and tip ;
outer surface of ears yellowish red ; mane light gray basally , fuscous
above; shoulder stripe absent; a dark but not very pronounced
median dorsal stripe extending from the lumbar region to the tail
tuft; forelegs yellowish gray anteriorly, light gray posteriorly; dark
bands about all the legs up to the level of the body, but only on the
anterior side of the forelegs (Noack, 1884, pp. 101-102). Height at
shoulder about 51 inches (Menges, 1887, p. 262). Pocock (1909,
p. 528) remarks on a seasonal change of color, from clear gray in
summer to sandy fawn in winter.
Menges (1887, pp. 263-267) gives the following account:
The range is apparently restricted to Somaliland and part of the
Red Sea coastal plain south of Massaua, Eritrea; it probably in-
cludes the Danakil region and extends south to the Webi Shebeli.
A particular habitat is the coastal lowlands, where the animal is
not exactly rare; another favored haunt is the barren Hekebo Pla-
teau (2,000 feet high), southeast of Bulhar. It is commonly found in
herds of 5 to 20 head, and is extremely shy and cautious.
In general it leads a rather undisturbed existence, though occa-
sionally falling victim to the Leopard or the Lion. Most of the
Somalis do not touch the flesh, but one or two tribes pursue the
animal to some extent. Only a few hides are brought from the
interior to the coastal markets. On the other hand, in the coast
districts of the Red Sea the Wild Ass is eagerly pursued, with the
object of shipping captured animals to Arabia, where they are used
for crossing with the Domestic Ass. One result of this crossing is
the hardy and beautiful riding ass of Yemen. Among the Somalis
themselves one finds many Domestic Asses bearing evidence of
crossing with the wild animals.
"In certain parts of Guban, notably in the sterile district lying
near the coast, about twenty miles east of Berbera, the Wild Ass is
not very uncommon. We met with it also in considerable numbers
on the high plateau west of Laferug, and also saw some individuals
south of the Golis Range .... The flesh of these animals is very
good, almost the best we ate in Somaliland .... It does not seem
to be a very plentiful species even in the country of its nativity,
and I should judge it would not require much persecution to speedily
extinguish the race." (Elliot, 1897, pp. 139-140.)
"The Somali wild ass is fairly common. I first met with them
about twenty miles to the south of Berbera, and they are also found
on the plateau to the south of the Golis range. They do not live on
the mountain ranges, but frequent the low stony hills in the desert.
352 EXTINCT AND VANISHING MAMMALS
They go in small herds. The largest I saw consisted of five." (In-
verarity, in Bryden, 1899, p. 71.)
''They are common to the eastward of Berbera, behind Siyaro, in
among the sand dunes and rocky hills, and also south of Bulhar in
similar localities, especially around the Issitugan Valley. South of
the Golis Range they inhabit the low stony hills around Halo,
Haloka Yer, and near Segig; they are also found on Negegr Plateau."
They are "usually seen in herds of four or five individuals and not
uncommonly singly." (Drake-Brockman, 1910, p. 103.)
Akeley (1914) gives an account of hunting Wild Asses about 30
miles from Berbera. One reason for their scarcity is indicated in
his statement (p. 117) that "one English 'sportsman' boasted of
having killed twenty-eight."
De Beaux (1928a, p. 6) records seven specimens from Italian
Danakil. He also remarks (p. 13) that the present subspecies is sepa-
rated from ajricanus by the Ethiopian plateau, which approaches the
Red Sea at the Gulf of Zula [Annesley Bay].
Thesiger, who traversed Abyssinian and Italian Danakil in 1933,
found Wild Asses quite common north and south of the lower
Hawash (Neumann, 1935, p. 153) .
Antonius (1938, p. 561) writes:
Because many hides from Berbera, as well as from Danakil, although typical
in all other points, show a more or less developed shoulder cross, there can
be no doubt that neither the existence of it nor its absence is thoroughly
typical [of somaliensis}. . . .
It is, alas! to be feared that the Abyssinian war has its consequences for
the African wild asses: warring soldiers, and especially askaris, are never
the best protectors of vanishing game! Whether the "Asinus somaliensis"
exists also in Southern Abyssinia or not is not positively known. A well-
informed Austrian, who had been living in Abyssinia for many years, told me
that he had seen wild asses in the Bale country on the upper Juba. I suggest
for geographical reasons that the animals are not true wild asses, but either
Zebras or domestic donkeys of a feral breed, similar to the beautiful asses
of the Turkana people on the western shore of Lake Rudolf, at first seen by
von Hoehnel and Count Teleky, and since recorded by modern visitors to that
country.
Lydekker (1916, vol. 5, p. 39) records a specimen from as far
south as "Shebeli Valley, Somaliland."
Atlas Wild Ass; Algerian Wild Ass
ASINUS ATLANTIOUS (Thomas)
Equus asinus atlanticus. P. Thomas, Mem. Soc. Ge"ol. France, ser. 3, vol. 3,
no. 2, p. 45, 1884. (Recent Quaternary deposits, Oued Seguen, near
Constantine, Algeria.)
SYNONYM: Equus asinus atlanticus Werth (1930).
FIGS.: Thomas, op. cit., pi. 2, figs. 7, 7a; Werth, 1930, p. 348, fig. 3; Jennison,
1937, pi. facing p. 145.
ORDER PERISSODACTYLA : ODD-TOED UNGULATES 353
Apparently a Wild Ass inhabited Algeria up to at least about
A. D. 300, but subsequently became extinct. It was probably distinct
from any form now living.
P. Thomas (1884, p. 45) based the name Equus asinus atlanticm
upon a mandible with teeth, found in late Quaternary deposits near
Constantine, Algeria. In these he found characters apparently inter-
mediate between those of Pliocene Hipparion and those of the
present-day Domestic Asses of Algeria.
The name atlanticus may be applied at least provisionally to the
Algerian Wild Ass of Roman times. Werth (1930, p. 350, map)
indicates the presumable former distribution as including Morocco
and Tunisia as well as Algeria.
"In a Roman villa at Bona, in Algeria, was found a large and
well-preserved picture, dating from about A. D. 300, of an African
hunt. Its main effect is a representation of a drive of carnivora. . . .
The use of the lasso is illustrated in the same picture, where a
Numidian, riding bareback and stirrupless, is throwing one at a wild
ass." (Jennison, 1937, pp. 145-146.)
Antonius (1938, pp. 559-560) says:
The true asses of African origin the wild stock from which our domestic
donkey descends belong to the many mammalia which became totally extinct
in our days. There were in Roman times at least three local races, one of
which became extinct before it was ever seen by a modern zoologist. It was
the "Asinus atlanticus Thomas," well known from the rock picture of Enfouss,
Algeria, published erroneously as "Quagga" by Frobenius. An excellent Roman
mosaic at Hippo Regius, the modern Bone, also shows that donkey. It
possessed a well-developed shoulder stripe, strongly marked limbs, and the
ears perhaps a little shorter than its East African cousins. The geographic
distribution of these Atlantic asses seems not to have exceeded the ranges
of the Atlas mountains. The time of their extinction is unknown.
Mongolian Wild Ass; Chigetai; Dziggetai; Kulan ; Kulon
ASINUS HEMIONUS HBMiONUS (Pallas)
Equus hemionus Pallas, Nov. Comm. Acad. Sci. Imper. Petropolitanae, vol. 19,
p. 394, pi. 7, 1775. ("Ad Lacum Tarei Davuriae" = Tarei Nor, on the
Siberian-Mongolian boundary, about lat. 50 N., long. 115 E.)
SYNONYMS: Equus onager castaneus Lydekker (1904); Equus (Asinus)
hemionus bedfordi Matschie (1911) ; Equus (Asinus) hemionus luteus
Matschie (1911). (C/. Harper, 1940, pp. 197-198.)
FIGS.: Pallas, op. cit., pi. 7, and 1781, pi. 1; Lydekker, 19046, pi. 27 (bedfordi) ;
Lydekker, 1904c, pi. 18 (castaneus); Lydekker, 1912, pi. 15, fig. 2
(castaneus); Lydekker, 1916, p. 13, fig. 6 (castaneus); Carruthers, 1913,
pis. facing pp. 602, 606; R. C. Andrews, 1924, pp. 152-156, figs., and 1926,
pi. facing p. 129.
During recent years, in all its vast range, the Mongolian Wild
Ass seems to have been reported as plentiful in only one region that
about Orok Nor and Zagan Nor in central Mongolia (about long.
354 EXTINCT AND VANISHING MAMMALS
100-102 E.). It has apparently disappeared from eastern Mon-
golia (including adjacent parts of Siberia and Manchuria). Wild
Asses throughout the world, with the apparent exception of the
Tibetan Kiang, are a vanishing type.
The following is derived from Pallas's early description (1781,
pp. 16-17) , supplemented by Radde's description (1862, pp. 293-294)
of what were virtually topotypical specimens from the vicinity of
Dalai Nor: The general color of the summer pelage is reddish yellow,
with a slight grayish tinge; in winter the color is more reddish than
yellow, and the hair is longer. Snout whitish ; rest of head more and
more yellowish ; mane brownish ; lower side of the neck of the general
body color; upper rump ochraceous; limbs and ventral surface paler
than sides; posterior side of forelegs, inner side of hind legs, lower
rump, and posterior border of the thighs whitish ; a brownish-black
median dorsal stripe from the mane to the bushy part of the tail,
broadest on the hindquarters; bristly hairs above the hoofs blackish.
Height at shoulder, about 3 feet 10 inches; length of ears, about 7
inches; length of tail without hairy tuft, about 1 foot 4 inches.
The former range apparently covered the greater part of Outer
Mongolia (except the present Tannu-Tuva) , small areas in Siberia
and Manchuria adjacent to the northeastern corner of Mongolia, at
least the western part of Inner Mongolia, and the northern part of
Chinese Turkestan (chiefly north of the Tian Shan) .
According to Pallas (1781, pp. 5-8), the Argun steppes are the
only place where these animals are still met with in Siberia. From
the rest of Dauria, where they once ranged, they have retreated into
the Mongolian deserts, on account of settlements. They still swarm
about Tarei Nor. Formerly they were seen on the Argun steppes in
great herds, but now only as solitary individuals or in scattered
troops. On the Mongolian Gobi they occur in numerous herds. This
is a game animal for the Mongols and Steppe Tungus, who eat the
flesh and make boots of the hide.
Radde writes (1862, p. 293) that in the fall and winter of 1856 a
strong northward migration extended to the region between Tarei
Nor and Dalai Nor, and that several animals were taken north of
Dalai Nor- (in northwestern Manchuria) . In a rare journal (Beitrage
Kenntniss Russ. Reiches, vol. 23, pp. 431-433) Radde gives addi-
tional information on life history, hunting, and economic uses.
A dearth of recent records of Wild Asses in eastern Mongolia
bodes ill for their survival in that region. They are evidently gone
from the adjacent parts of Siberia and Manchuria. Arthur de C.
Sowerby (in Hit., March 14, 1938) believes they have ceased to
exist in all these areas.
In 1887, in the region about the southern base of the eastern Altai,
ORDER PERISSODACTYLA I ODD-TOED UNGULATES 355
at about longitude 96-100 E., Younghusband (1888, p. 495) saw
"considerable numbers of wild asses."
In the central Gobi, in 1922-25, R. C. Andrews (1924, pp. 152-154;
1926, pp. 132-145, 299-302, 317-318) found considerable numbers
of Wild Asses in the vicinity of Orok Nor and Zagan Nor (about
long. 100-102 E.) "During the first two years of our work in the
Gobi, we never saw wild asses in herds of more than fifteen or
twenty, but we did not arrive in their country until after the breed-
ing-season. In 1925 the herds numbered thousands. Evidently
they collect at favorable localities just before the young are born .
. . . The young are dropped about the beginning of July, and the
asses seek a flat plain, undoubtedly for protection from wolves."
(P. 302.)
"I have been asked by many people if it would be possible to
catch wild asses when they are young and use them for breeding
purposes. I do not believe that this would be practicable, due to
the extraordinary wildness of the animals. Certainly, it would be
difficult to tame an adult wild ass." (R. C. Andrews, 1924, p. 154.)
In 1926 the Kulan was very common at the northern base of Iche-
Bogdo, in the valley of the lakes west of Orok Nor, and in the desert
area to the northwest as far as the Baidarik River (Formozow,
1931, p. 77).
In 1911 Carruthers (1913, p. 532) found that the western shore
of Bar Kul, in southern Dzungaria, was the haunt of droves of
Wild Asses. His companion, J. H. Miller, supplies much additional
information (in Carruthers, 1913, pp. 582, 588-589, 603-608). In
the hills west of Bar Kul, towards the end of April, "a few wild-
asses, straight from their winter quarters on the lowlands to the
north, were busy making up for their scanty winter fare" (p. 588) .
In the vicinity of Shi-Kho, at the northern base of the Tian Shan,
a domesticated Kulon was examined; it was perfectly docile, but
could not be broken to the saddle. A large wild herd was seen in the
same area. (P. 603.) In Guchen a Kirghiz reported Kulon very
numerous in the sand-dune area to the north, and Miller himself
found fair numbers there (p. 604) . Two specimens were secured near
the Dzungarian Gate north of Ebi Nor, where a spring was much fre-
quented by Kulon (p. 605) .
"My specimens are undoubtedly Equus hemionus typicus ....
Its extreme eastern distribution is at present imperfectly known;
Sir Francis Younghusband, in his journey across the Northern Gobi,
mentions seeing kulon in the Gobi at the extreme eastern end of the
Altai. They are found north of the Altai Range on the plains,
round the large lakes in the Kobdo region; we met with them near
Barkul, and in several other places throughout Southern Dzungaria.
. . . The natives hunt them occasionally for their skins and meat,
356 EXTINCT AND VANISHING MAMMALS
which they consider more palatable that the best mutton." (J. H.
Miller, in Carruthers, 1913, pp. 607-608.)
"The wild ass, or kulon, is unlikely to be seen unless a special
attempt is made. . . . The kulon is a rare animal, excessively wild
and lives in very difficult country. Featureless plains, bitterly cold
in winter, waterless and sunbaked in summer, are its habitat. The
kulon ranges . . . through Dzungaria to the edge of the Gobi. We
have seen them at the lowest elevation in the heart of the conti-
nent, and at 7,000 to 8,000 feet above the sea, in localities not very
far distant from each other." (Carruthers, 1915, p. 154.)
In 1926 a journey made by Lattimore (1929, pp. 228-321)
through the southwestern Gobi filled in some blank spaces in the
known distribution of this species. West of Edsin Gol, at the Wild
Horse Well (about lat. 42 N., long. 99 E.) : "They say that on
this fringe of the Khara Gobi there are wild horses . . . and wild
asses" (p. 228). Near the "House of the False Lama" (about lat,
42 30' N., long. 98 E.) : "To this whole series of springs there
come at night antelope, wild asses, and, they say, wild camels" (p.
243) . In the vicinity of Ming Shui (about lat. 43 N., long. 96 E.) :
"To our camp that day there came riding a Mongol, who had fol-
lowed us for two marches to sell the hinder half of a wild ass that
he had shot in the Mongol way from a pit near the drinking place"
(p. 251) . West of Ming Shui, near the eastern outposts of the Kar-
lik Tagh: "Here the camel herders in the dawn reported a herd
of wild asses. It was the only sight of them that I ever had ....
Their skins make first-class clothing, with much more wear than the
antelope skin. I have heard that there is a Turki proverb that wild
asses are so hard to kill that even when you get the skin of one
safely spread out on your sleeping platform it wiggles. The meat
is something like beef, but a sublime beef. It is very dry, with a
coarse grain and a strange aromatic sweetness. Chinese and Mon-
gols put it above any other game, and it undoubtedly ranks with
the noblest vension." (P. 252.) At Wu-t'ung Wo-tze, about 100
miles northeast of Kucheng, in the Dzungarian Gobi: "It . . . was
formerly a well-known wild-ass ground; but the wild ass in this
region has been almost killed off by the Qazaqs. Both Mongols and
Qazaqs will put themselves to more trouble to bag wild ass than
almost any other game." (P. 321.)
Farther southwest than the territory covered by Lattimore, along
the route from Hami to Bulundsir River, Wild Asses were
reported in 1898 in a number of places by Futterer (1901, pp. 179,
180, 184, 188) . A specimen obtained northwest of the last-mentioned
locality became the type of Equus hemionus luteus Matschie.
In 1934 Sven Hedin (1940, pp. 195, 197, 200) found tracks in the
Ghashun Gobi about 75 miles west of Futterer 's route. This area
ORDER PERISSODACTYLA : ODD-TOED UNGULATES 357
seems to constitute the southwestern limit (as far as known at
present) of the range of the Mongolian Wild Ass.
Enemies. Among predatory animals, the Wolf seems to be the
only enemy of any importance, and doubtless it has never affected
the Wild Asses at all seriously. Apparently it cannot successfully
attack any except the young Asses within a few weeks of their
birth. Older animals are able to outrun the Wolf on the open plains.
Increasing use and precision of firearms in the hands of the
Asiatics have undoubtedly contributed chiefly to the decline of the
AVild Asses.
Transcaspian Wild Ass; Transcaspian Kulan; Wild Ass of
Russian Turkestan. Transkaspischer Kulan (Ger.)
ASINUS HEMIONUS FiNscHi (Matschie)
Equus (Asinus) hemionus finschi Matschie, in Futterer, Durch Asien,
vol. 3, pt. 5, Zoologie (Nachtrag), p. 24, 1911. ("Nordostlich vom Saisan-
nor" (Zaisan Nor, in former Province of Semipalatinsk, Russian Turke-
stan).) (Cf. Harper, 1940, p. 198.)
FIGS.: Radde, Sammlungen Kaukas. Mus., vol. 1, Zoologie, pi. facing p. 60,
1899; Brehm's Tierleben (IV), 12, p. 670, tab. Unpaarhufer V, fig. 2,
1915; Schwarz, 1929, p. 92, fig. 5.
This Wild Ass is now very scarce in Russian Turkestan, having
evidently disappeared from the greater part of the country. It was
long ago exterminated in southern Russia.
Matschie describes the type (from the vicinity of Zaisan Nor)
as reddish salmon, with a slight tinge of gray; the lips are white;
the whitish of the under parts extends well up on the flanks; the
dark vertebral stripe continues on to the base of the tail. Schwarz
adds (1929, p. 91) that the maximum width of the vertebral stripe
is 42 mm. Radde and Walter (1889, p. 1059) describe a full-grown
male from the Askabad region as lacking a shoulder stripe; its
height at the shoulder was 1,110 mm.; tail (including tuft) , 590 mm.
"It is clear [from Strabo's account] that the wild ass (onager)
existed all across southern Russia in the fifth century B. C., for it
was hunted both by the Sarmatian tribes who lived on the east side
of the Don (Tanais) and by the Scythians who occupied the region
to the west of that river. It is even possible that the wild ass dwelt
in the Danube valley almost down to the beginning of the historical
period. It seems certain that neither Sarmatian nor Scythian ever
domesticated the wild ass, a circumstance probably due to the fact
that they had a more docile and serviceable animal in the wild
horses of the same region." (Ridgeway, 1905, pp. 51-52.)
"In former days kulan and onagers appear to have ranged much
further westward than is the case at the present day. It is stated,
for instance, by the Russian naturalist Rytschkov that in the
eighteenth century kulan abounded on the eastern side of the Volga,
358 EXTINCT AND VANISHING MAMMALS
and from time to time troops swam that river and made their ap-
pearance in the Waldinsel Steppe." (Lydekker, 1912, p. 185.)
Pallas gives the following account in 1780 (pp. 261, 273). Kulans
are still very numerous in the deserts of Tatary, and come annually
to spread over the mountainous deserts east and north of the Aral
Sea, where they pass the summer and assemble in autumn by hun-
dreds and even thousands for their return toward India, for a winter
asylum. They scarcely pass beyond lat. 48 N. The skins are
sought by the residents of Bokhara for preparation in the manner
of shagreen.
Hablizl states (1783, p. 93) that the Wild Ass is rarely found in
Astrabad and Mazanderan, Persia. (The animal of this Caspian
region was probably the Turkestan form rather than the Persian.)
In 1840 (p. 56) Eversmmann records several recent specimens
from the steppes between the Caspian and the Aral Seas.
Ladyjensky (1841, pp. 361-362) speaks of sending to Moscow a
Wild Ass captured when young in the vicinity of Aktava in the
Kirghiz Steppe, in the extreme southern part of the old district of
Omsk. In this region, he adds, the Wild Asses are found in numerous
troops, which are composed sometimes of more than a thousand
head and raise a thick cloud of dust in moving across the steppe.
The Kirghiz have not yet found the means of taking the young ones
alive.
Severtzoff writes (1876, p. 387) that Equus hemionus "is rather
rare in Turkestan, and to be found only about the Karatau moun-
tains and near the rivers Aris, Keless, Chirchik, and the delta of the
Sir-Darja, and even there only during the winter."
In 1881 (p. 22) Poliakof refers to the Kulan as having been numer-
ous not long since in the environs of Lake Balkash. "In my last
excursion to Balkash, during several days passed in its solitudes I
did not observe a kulan, and only saw the tracks of one imprinted
on the saline soil."
According to Radde and Walter (1889, p. 1058) , it still inhabited
at that time the entire Turkoman Steppes in considerable numbers,
but had retreated from the Transcaspian railway and the new mili-
tary posts farther into the undisturbed deserts. At the beginning of
the construction large herds were often observed near the railway in
the vicinity of Kazanjik and Dushakh, but later disappeared. It
was considered more numerous in the steppes north of the Atrek
River and also along the Afghanistan boundary in the hilly desert
between the Tejend and the Murghab. Here Walter saw many in
1887. While the European hunter rarely had success, the Saryk
Turkoman managed to approach within gun range of the animals by
taking cover behind a carefully maneuvered camel. The flesh was
ORDER PERISSODACTYLA : ODD-TOED UNGULATES 359
much prized by the Turkomans, and in winter was commonly sold
at the bazaar in Yolatan.
Matschie (1911, p. 23), besides recording the type of finschi from
northeast of Zaisan Nor, mentions another specimen from the
Maiterek Steppe north of this lake.
J. H. Miller (in Carruthers, 1913, p. 608) states that the animals
" extend throughout Northern Russian Turkestan, being exceedingly
numerous in the neighbourhood of Lake Balkash." It is doubtful,
however, if this statement was appropriate as late as 1913.
Schwarz (1929, p. 91) mentions a specimen from the vicinity of
Merv that formerly lived in the Berlin Zoological Garden. He gives
the range of this form as extending from the northern border of the
Persian Plateau through West Turkestan and the Kirghiz Steppe to
the western slope of the Altai.
Nazaroff (1932, p. 54) refers to the species as extinct in the Tash-
kent region.
According to W. G. Heptner (in litt., December, 1936) , the Kulan
is now a rare animal with a small distribution in the U. S. S. R.
At the beginning of the nineteenth century it occupied the plains of
Turkestan and almost all the steppes of Kazakstan. It is now met
with in Turkmenia (principally west of the Murghab) , and persists
in small numbers in the desert steppes near Lake Balkash. Hunting
is absolutely forbidden, and reserves are being organized.
"Today there are only a few scattered troops, in yearly diminished
numbers, near the Oasis of Merw and the Afghanistan frontier the
last remnants of the immense herds which roamed the steppes of
Asiatic Russia 100 years ago" (Antonius, 1938, p. 559) .
North Persian Wild Ass; Persian Onager; Ghor-khar
ASINUS HEMIONUS ONAGER (Boddaert)
[Equus] Onager Boddaert, Elenchus Animalium, p. 160, (1784) 1785. (Based
upon the "Onager" of Pallas, Neue Nord. Beytrage, vol. 2, p. 22, pi. 2,
1781; type locality erroneously stated as "in desertis Argunis"; corrected
by Harper (1940, p. 199) to "mountains about Kasbin, " northwestern
Persia.)
FIGS.: Pallas, 1780, pis. 11-12; Pallas, 1781, pi. 2; Pallas, Zoographia Rosso-
Asiatica, pi. to vol. 1, p. 264, 1834-42; Hamilton Smith, 1841, pi. 18;
Lydekker, 1904, pi. 19; Kennion, 1911, pi. facing p. 121; Brehm's Tier-
leben (IV), 12, p. 674, tab. Unpaarhufer V, fig. 3, 1915; Antonius, 1939,
figs. 2-3.
The present numerical status of the North Persian Wild Ass,
like that of various other mammals of that country, is a matter on
which up-to-date information is difficult to acquire. It is probably
quite scarce; at least it very seldom affords a glimpse of itself to
the scientific traveler. The Wild Ass of Afghanistan is provisionally
referred to the present subspecies.
360 EXTINCT AND VANISHING MAMMALS
Both Pallas (1781, pp. 34-37) and Hablizl (1783, pp. 89-92)
furnished descriptions of the original two specimens (male and
female) from Kasbin. Since Hablizl alone was acquainted with the
male in life, his description of that sex may be considered the more
dependable, and is here utilized. In the male the top and sides of
the head are half-reddish, the lower side and the muzzle white;
outer surface of ears reddish yellow at base and tip, white in the
middle; sides of neck and of body and fore part of haunch reddish
yellow (isabelline) ; mane light brown, composed of hairs 3-4 inches
long; a light-brown vertebral stripe, up to 2^ to 3 inches wide, extend-
ing from the mane to end of the tail; a similar but smaller stripe
crossing this at right angles on the shoulders; lower neck, lower
shoulder, breast, belly, buttocks, vertical area in front of thigh,
dorsal area bordering the vertebral stripe, and legs white; tail like
a cow's, with a tuft of long, light-brown and white hairs. Height at
shoulder, 50 inches; ear, 11^ inches; tail (including tuft), 25 inches.
The female is similar, but smaller, and lacks the shoulder stripe.
Height at shoulder, 44 inches; ear, 8J inches; tail, 20 inches.
The three skins from Yezd, Persia, listed by Lydekker (1916,
vol. 5, pp. 14-15) , are very white on the sides and belly and have no
shoulder stripe (J. C. Phillips, in litt., June, 1938). Whether these
specimens are all females, or whether some males lack the shoulder
stripe, is difficult to say. Goodwin (1940, p. 17) decribes the summer
pelage as either avellaneous or light pinkish cinnamon, with a faint
shoulder stripe ; and the winter pelage as sayal brown, without a
shoulder stripe.
Persia. The Persian range of this Wild Ass is here considered
restricted to a portion of the Persian Plateau, extending north to the
Elburz Mountains, east to Afghanistan, south to about the latitude
of Seistan, Kerman, and Shiraz, and west (at least formerly) to the
mountain ranges extending along the line Kasbin-Ispahan-Shiraz. 1
Since Omar Khayyam was a resident of Naishapur in Khorassan,
it was doubtless the present subspecies that he had in mind in the
following verse:
They say the Lion and the Lizard keep
The Courts where Jamshyd gloried and drank deep :
And Bahrain, that great Hunter the Wild Ass
Stamps o'er his head, but cannot break his Sleep.
1 There is considerable uncertainty, however, as to the boundary between the
ranges of this and the Indian subspecies. For example, Lydekker (1904, p.
589, pi. 17) refers to the latter a male in the London Zoo that was said
(probably erroneously) to have come from the desert near Meshed, in north-
eastern Persia; it lacked the shoulder stripe generally considered diagnostic
of onager. There seem to be extremely few records of specimens with shoulder
stripes and from a definite locality.
ORDER PERISSODACTYLA ! ODD-TOED UNGULATES 361
According to Pallas (1780, pp. 259-273), Onagers are found in
the mountains about Kasbin at all times of the year. The Persians
seek to take them alive in pitfalls, driving them toward these spots.
The young ones captured alive are sold for a considerable price for
the studs of the nobles. From the mating of these tame Onagers there
is derived the fine race of riding asses in Persia and Arabia. They
are clearly distinguished from the sorry race of ordinary asses that
are used for carrying burdens. The bile is esteemed among the
Persians as a remedy for obscure vision and for cataract.
Hablizl writes (1783, pp. 93-94) that the Wild Ass is frequently
seen in herds in the valleys of the mountains about Kasbin. Yet it
is reckoned among the rarest animals in all Persia, being known to
most of the inhabitants only by name. It is captured only to be
presented to some Khan, who has it kept as a mere rarity. In a few
months it becomes completely tame, more especially if it has been
caught while young.
"In the eastern provinces of Persia . . . their venison is highly
prized, and the chase of them, from the time of Rustum to the
present, has always been held the pastime of heroes and princes"
(Hamilton Smith, 1841, pp. 309-310) .
Blanford writes (1876, p. 85) : "In Persia they appear ... to be
common in some places, generally on the borders of desert plains,
rare or unknown elsewhere; but they occur scattered over all the
more level parts of the country, except in the North-western and
Caspian provinces. I saw none during my journey, though I often
came across their tracks."
To this St. John adds (in Blanford, 1876, pp. 85-86) : "All of
twenty or more specimens that I have seen from Western Persia were
undoubtedly E. onager .... The Persian wild ass is not, I
believe, found west of the main road from Tehran to Shiraz, except
possibly in that arm of the salt desert which extends north of Korn-
. . . towards Saveh. It is most plentiful in the vicinity of the
patches of salt desert, 'Kafah' or 'Kavir/ which are so marked a
character of Eastern Persia. In the summer a herd occasionally
wanders into the loftier desert valleys. I have several times seen
them whilst travelling post along the plain that stretches from
Khan-i-Khora, a short distance north of Dehbid, in Fars, to the
Kulah Kazi or Urchin Hills, near Isfahan, a distance of nearly 150
miles, at an elevation of 5500 to 7000 feet above the sea. Persians
say that they can not be caught by a single horseman when ap-
proached in the open; but if the sportsman can manage to conceal
himself and his horse in the vicinity of a spring, and wait until the
wild asses have quenched their thirst, they can readily be come up
with when full of water, by a short spurt on a fast horse. At other
times they are caught in relays of horsemen and greyhounds. The
\
362 EXTINCT AND VANISHING MAMMALS
flesh is said in books on Persia to be prized above all other venison ;
but Persians have told me that it should only be eaten under absolute
necessity, being equally disagreeable to the conscience of a good
Mussulman, and to the palate of a gourmand."
In his journey of 1906, Sven Hedin (1910, vol. 1, pp. 216, 218, 222,
228, 243-244, 396-397; vol. 2, p. 65) saw a few Wild Asses and
noted tracks or received reports of numerous others on the western
and southern borders of the Great Salt Desert ("Kevir") in north-
eastern Persia. On the western border "the herdsmen . . . are ac-
customed to the presence of wild asses, and take no notice of them."
On the southern border a native "had shot many wild asses, and was
wont to sell their skins to the shoemakers of Tebbes. When, as now,
there was much rain, the chase was not profitable, for the wild asses
could find water anywhere; at other times they are dependent on
springs, and then is the time for the huntsman to stalk his prey."
Farther south, in Kuhistan, the animals were said to be very
numerous in the desert between Tebbes and Bahabad, and a local
hunter "had killed two hundred wild asses."
"Most of the remoter deserts of Eastern Persia are roamed over
by wild asses. Unfortunately, . . . these animals have always been
remorselessly persecuted for the sake of their meat and hides,
usually by the ignoble plan of sitting up over water, with the con-
sequence that they are now scarce." (Kennion, 1911, p. 119.)
"His [the Anatolian Onager's] near ally in Iran (Equus hemionus
onager Zimm.) , although still rambling over the salt deserts of his
country in considerable numbers, is hard pressed by modern guns"
(Antonius, 1938, p. 559) .
The most recent account is by Legendre (1939, pp. 240-241) :
"Goodwin had succeeded in shooting one, 50 miles from Teheran.
"At the village of Abbasabad [north of the Great Salt Desert]
we received information that there were herds of wild ass to be
found in the desert 80 miles away, near a salt spring." At this
spring "there were tracks everywhere," but none of the animals could
be located.
Two natives reported "that the wild asses had all migrated to
the south two months before .... They assured us, however,
that they were to be found in herds of ten to forty around the salt
spring, from June until September."
The locality referred to by Legendre is evidently identified by
Goodwin (1940, p. 17) as Siah Parde, whence he records three
specimens.
Afghanistan. Scarcely any information seems to be available
concerning the Wild Ass in this country, save that secured by the
ORDER PERISSODACTYLA : ODD-TOED UNGULATES 363
Afghan Delimitation Commission of 1884-85 (Aitchison, 1889,
pp. 61-62) :
[On November 30, 1884, herds were seen on the march between Tut-i-chi
and Aftao, north of the Paropamisus Range.] They occupied the country
in the vicinity of Gulran, as they were known to have attacked and injured
some Mules and Donkeys that had been turned loose to graze. . . .
In my march from Gal-i-cha [south of the Paropamisus Range] to the
base of the Kambao Pass, on the 29th of April, 1885, I had to cross the
northern end of a great plain called "Gulam-i-maidan," or the plain of the Wild
Ass. . . . My guide took me to a slight elevation, and from it pointed
out to me where I was to look for the animals: for some time I could see
nothing ; at last, whilst using my glasses, I noticed clouds of dust, like the line
of smoke left in the track of steamers. . . . These several lines of dust-cloud
were caused by herds of Asses, galloping in various directions over the great
plain. One herd came well within a mile's distance ; from its extent, I am
even now of the opinion which I then held, that the herd consisted of at
least 1000 animals. I counted sixteen of these lines of dust-cloud at one
time on the horizon. My guide said that at this period of the year the Wild
Asses are always united in great herds on that plain, owing to the mothers
having their foals at foot, but that in a few weeks the great herds would
break up, and the animals would spread themselves all over the country in
parties of ten to twelve. This is the season at which the young are caught,
by riding them down; usually, the mother will not leave, viciously attacking
men and horses upon their coming near her foal. It is a very rare circum-
stance to get a foal unless by shooting it. ...
Between Karez-dasht and Sher-baksh [lat. 33-34 N.], to the south-east of
the Do Shakh range, we were informed was a locality for the Wild Ass,
also the country between Kushk-rud and Zagin, still further south, but I did
not hear of any having been seen by members of the Mission.
It is undoubtedly too much to hope that Wild Asses abound in
such numbers today, as they did half a century ago, in north-
western Afghanistan.
Indian Wild Ass; Baluchi Wild Ass; Ghor-khar. Ane de Hnde
(Fr.). Indischer Wildesel (Ger.).
ASINUS HEMIONUS KHUR (LeSSOn)
Equus khur Lesson, Manuel Mammalogie, p. 347, 1827. (Based upon "der
wilde Esel" of Oken's Isis, 1823, Band 2, Heft 7, p. 764, 1823, inhabiting
"die wiiste Strecke Landes, welche Cattuwar von Cuth trennt (bey den
Eingebornen Run genannt)." Thus the Little Rann of Cutch, western
India, is the type locality.)
SYNONYMS: Equus indicus George (1869) ; Equus hemionus var. indicus W. L.
Sclater (1891).
FIGS.: Cuvier, Regne Animal, disciples' ed., Mamm., atlas, pi. 83, fig. 1,
1836-49; J. E. Gray, Gleanings from Knowsley Menagerie, pi. 53, 1850;
Lydekker, 1904, pi. 17 (ssp.?); N. Y. Zool. Soc. Bull., vol. 24, no. 1,
p. 12, fig., 1921; Schwarz, 1929, figs. 1-4; Jour. Bombay Nat. Hist. Soc.,
vol. 37, no. 1, suppl., pi. 29, 1934.
The range limits of this subspecies toward the north and west are
uncertain. The Wild Ass of southeastern Persia will be provisionally
364 EXTINCT AND VANISHING MAMMALS
considered the same form as that of Baluchistan and western India.
The animal has evidently disappeared over a great part of this
range. It survives in small numbers in the Rann of Cutch, India,
and probably in some of the desert tracts of southeastern Persia.
The general color (in summer) is sandy ; muzzle, buttocks, breast,
lower parts, and inside of limbs white x ; ears sandy externally, white
internally, with a blackish tip and outer border; mane and tail tuft
blackish brown; a chocolate-brown vertebral stripe extending from
mane to tail, bordered from the withers backward by a light area;
front of all four limbs very light, with a slight yellowish tinge; a
narrow blackish ring above the hoofs. The winter pelage is longer,
and grayish. Height of male at shoulder, about 47 inches. (Chiefly
from Jerdon, 1874, pp. 236-237, and Schwarz, 1929, pp. 87-88.) An
adult female from the Punjab-Sind frontier measured: height, 46
inches; tail (including hair), 26 inches; ear from crown, 9 inches
(Blanford, 1891, p. 470).
The absence of a shoulder stripe in the male, the presence of a
blackish ring above the hoofs, and apparently the less pure white
of the lower parts, may serve to distinguish the Indian from the
North Persian subspecies (onager).
The place of this animal in ancient history is sketched by Ridge-
way (1905, pp. 47-48). According to Herodotus (VII, 86), "some
of the Indians in the army of Xerxes drove chariots drawn by 'wild
asses.'
"From this it is clear that the peoples of western Hindustan, who
did not possess horses, had made the wild ass obedient to the yoke.
"In Carmania . . . , a region bounded by the Indian Ocean and
Persian Gulf on the south, and by Persia on the west, down to the
time of Strabo, 'asses on account of the scarcity of horses' were
'generally made use of in war. They sacrifice an ass to Ares, who is
the only god worshipped by them, for they are a warlike people.' "
According to an anonymous writer in Oken's Isis (Band 2, Heft
7, p. 764, 1823) , herds numbering up to 60 or 70 were observed in the
Rann of Cutch. The animals are said by the natives to be very shy
and hard to capture. In November and December they come deeper
into the land, in herds of hundreds, and cause great damage in the
cultivated fields. Therefore they are caught in pitfalls. The flesh
is considered good by many people of the lower classes, who lie in
wait for them when they come to drink.
The breeding of captive animals from Hindustan was successfully
carried out in Paris from 1842 to 1849. Of nine foals produced
during this period, six survived in 1849. The animal was also said
i A specimen recorded by Lydekker (1916, vol. 5, p. 13) from the Sham Plains.
Baluchistan, is not so pure white on the lighter parts as three specimens of
A. h. onager from Yezd, Persia (J. C. Phillips, in Hit., June, 1938).
ORDER PERISSODACTYLA : ODD-TOED UNGULATES
365
to be used occasionally in its native land for agricultural work.
(I. Geoffroy, 1849, p. 35.)
"The ghorkhur is found sparingly in Cutch, Guzrat, Jeysalmeer
and Bikaneer, not being found further south, it is said, than Deesa,
or east of 75 east longitude. It also occurs in Sindh, and more
abundantly west of the Indus river, in Beluchistan .... It appears
that the Bikaneer herd consists at most of about 150 individuals
. . . . " A writer in the Indian Sporting Review is quoted on the
FIG. 38. Indian Wild Ass (Asinus hemionus khur)
animal's occurrence in the desert country west of the Indus, above
Mithunkote : "The foaling season is in June, July, and August,
when the Beluchis ride down and catch numbers of foals, finding
a ready sale in the cantonments for them, as they are taken down on
speculation to Hindustan. They also shoot great numbers of full-
grown ones for food, the ground in places in the desert being very
favourable for stalking." The same method of capturing foals is
practiced in Bikaneer. (Jerdon, 1874, p. 238.)
"Wild asses are renowned for speed, but in the Rann of Cutch
adults have been run down by men on horseback and speared. I
believe, however, the animals run down were mares in foal." (Blan-
ford, 1891, p. 471.)
"I told his Highness of Bikaner in London last autumn that I
wished to intercede with him 'on behalf of an oppressed minority
366 EXTINCT AND VANISHING MAMMALS
resident in his dominions'; the official books allege that Bikaner has
'a herd of about 100 wild asses/ He assured me the books were
wrong; he himself had never seen one, but very occasionally one
strayed into his borders from Bhawalpur. They still exist in Kutch,
or did until yesterday. These too are in an isolated area where they
cannot be replenished from outside." (Edward Thompson, in Lon-
don Times, August 19, 1932?)
"Within recent years, it has become confined to the Rann of
Kutch .... The depletion in numbers is attributed to -the fact
that it has for long been hunted by certain tribes for food; and it
is significant that it now only exists in appreciable numbers in an
inaccessible locality like the Rann, where it is rigorously protected
by His Highness' Government. The killing ... is illegal throughout
His Highness' territory; and such killing as at present takes place is
due entirely to hunters from British India and from the States on the
Indian side of the Rann, who occasionally make raids upon the
animal, either to secure a rare trophy or to kill it for food. This
Government contemplates addressing a request to the British author-
ities that the killing of the animal should be made an offence else-
where, on the analogy of the rules already in force in Kutch. But the
wild Asses found within Kutch jurisdiction, and not straying outside
it, are perfectly secure." (Dewan's Office, Kutch, in litt., February,
1937).
The present range in India is the eastern parts of the Rann of
Cutch, in very reduced numbers. The Wild Ass has been recom-
mended by the All-India Conference for the Protection of Wild Life
as one of the species which should be specially protected. The testes
are believed to possess aphrodisiac properties. (Bombay Natural
History Society, in litt., December, 1936) .
Blanford says (1876, p. 85) : "Wild asses are locally distributed
in [Persian] Baluchistan, and I only heard of their being abundant
near Bampur. None are said to be found in the deserts north of
Jalk and Kalagan [in northwestern Baluchistan], though Ferrier
speaks of them as common farther north in Sistan." Detailed
reports from Baluchistan of more recent date do not seem to be
available.
Lydekker (1904, p. 589, pi. 17) records a male specimen in the
London Zoo, apparently referable to this subspecies, and "stated
to have been captured as a foal in the desert near Meshed," north-
eastern Persia. The alleged provenance, however, may be regarded
as distinctly questionable. Schwarz (1929, p. 91) refers this speci-
men to onager, despite the impure white of the lower parts and the
lack of a shoulder stripe.
Schwarz (1929, pp. 85-89, figs. 1-4) describes and figures a
specimen in the Berlin Zoological Garden, evidently belonging to
ORDER PERISSODACTYLA: ODD-TOED UNGULATES 367
the Indian subspecies and said to have come from the desert north-
east of Kerman, Persia. He regards this locality as the present
western limit for the subspecies, although he makes Asinus hamar
Hamilton Smith (from the northern part of the Province of Fars)
a synonym of A. h. khur. I prefer to consider hamar indeterminable
at the present time.
Syrian Wild Ass. Hemippe de Syrie (Fr.) Syrischer
Halbesel (Ger.)
ASINUS HEMIONUS HEMIPPUS (I. Geoffrey)
Equus hemippus I. Geoff roy-Saint-Hilaire, C. R. Acad. Sci. [Paris], vol. 41,
p. 1214, (1855) 1856. (Based upon two live captives, said to have come
"du desert de Syrie, entre Palmyre et Bagdad" (op. cit., p. 1219,
footnote).)
FIGS.: Milne-Edwards, Nouv. Archives Mus. Hist. Nat. Paris, vol. 5, Bulletin,
pi. 4, 1869 (cotype); Antonius, 1928, figs. 1-5, and 1939, fig. 1.
The question of the Wild Asses of the Syria-Palestine-Arabia-
Iraq region is a very troublesome one, especially on account of the
meagerness of material and information. It is further complicated
by the fact that a number of authors recognize two distinct species
in this region, although in no other part of the world are as many
as two different forms of Wild Asses definitely known to occur to-
gether. The Wild Ass of this region must be on the verge of extinc-
tion, if not already extinct.
Since Geoffrey's cotypes (1856, p. 1217) were both subadult
females, we shall turn to Antonius (1928, pp. 21-22) for a description
of both sexes of hemippus. This is the smallest form of Recent
Equidae. The general color of the male is "avellaneous" (Ridgway) ,
becoming a sort of mouse gray with age ; the color is lightest on the
head, darkest on the haunches; a light area in front of hips; buttocks,
belly, and inner side of legs dirty grayish white; outer side of legs,
lower side of neck, and outer surface of ears "tilleul buff"; tips of
ears originally dark brown, later almost white; mane rather long,
"natal brown"; vertebral stripe, of the same color, extending from
the mane to the tail tuft, and bordered by a lighter area; area
above the nostrils grayish white; nostrils very large and nasal
region swollen. Height at shoulder, 1 meter.
The general color of the female is between avellaneous and fawn
color; buttocks and lower parts pure white; outer side of legs and
ears "pinkish buff"; tips of latter scarcely darker. Height at
shoulder, 1 meter.
Tristram (1884), Aharoni (1930), and Bodenheimer (1935) fail
to assign any adequate diagnostic characters to the two kinds of
Wild Asses that they recognize; they also place both of them in the
368 EXTINCT AND VANISHING MAMMALS
Syrian Desert and neighboring regions. Thus the two forms must be
specifically distinct, if distinct at all. The names applied are onager
and hem,ippus, which are currently considered subspecies of a single
species. It is almost unquestionable that the name Asinus hemionus
onager (Boddaert) , typified by the Wild Ass of northwestern Persia,
cannot be applied to a form of the Syrian Desert. If there was a
Wild Ass in this region distinct from hemippus, some other name than
onager must be found for it.
In view of the extension of various African types of mammals
past the Isthmus of Suez into Syria and Arabia, it is perhaps not
beyond the bounds of possibility that, if there was a second form of
Wild Ass in the Syrian Desert, it was some form of the African
Wild Ass (A sinus asinus) . A character this species has in common
with the Persian Onager is a shoulder stripe. The type of asinus,
according to Lydekker (1916, p. 37), is the domesticated ass of
Asia; but he also remarks (1912, p. 217) that we have no evidence
that its wild progenitor ever existed to the eastward of the Red Sea.
A different view is expressed by Tristram and by Ridgeway, who are
quoted below. It is perhaps now too late to secure conclusive light
on the subject. The following account will include both alleged
forms.
Tristram says (1884, pp. 2-3) concerning "Asinus onager": "This
Wild Ass, the origin of the Domestic Ass, was formerly well known
in Arabia, and is not extinct there, though very rare. I have seen
this species in a state of nature frequently in the Sahara, and have
handled captured though not tamed individuals. It no doubt, as the
Arabs assure me, occasionally enters the Hauran [at the north of
the Syrian Desert]. Their language, as well as the Hebrew, recog-
nises two species of Wild Ass."
Ridgeway (1905, pp. 52-53) writes in similar vein: "There is
strong evidence that the Arabs had domesticated some kind of E.
hemionus from a very early time, for we shall find later on that the
Arab tribes possessed asses from the dawn of history, and Strabo
when describing the littoral of the Red Sea after Eratosthenes and
Artemidorus, speaks of a region south of Nabataea well wooded and
well watered, abounding with all kinds of cattle, wild asses
(hemionoi), wild camels, deer, and gazelles .... As there were
thus both wild asses and wild camels in Arabia down to the Christian
era, there can be little doubt that the domestic asses and camels of
the Arab tribes were derived from the wild species of that region."
Tristram also gives (1884, p. 3) the following account of "Asinus
hemippus": "This, rather smaller than the true Onager, and con-
fined to Syria, Mesopotamia and North Arabia, very rarely enters
the north of Palestine from the Syrian desert, but is still common
in Mesopotamia. It does not extend into India, but in summer herds
ORDER PERISSODACTYLA I ODD-TOED UNGULATES 369
of this animal frequently visit the Armenian mountains. It is the
Wild Ass of Scripture and of the Ninevite sculptures."
"The fluctuations of the Beduin troops during the World War in
general and of the Wahabi tribes in late years have quite wiped out
the limits of both species of Wild Asses (Asinus hemippus and A.
onager) reaching Trans-Jordania, and have pushed back these
extraordinarily shy, freedom-loving animals into the center of the
desert. They now occur so sporadically that many Beduin clans
have not seen them at all during late years." (Aharoni, 1930, p. 330;
translation.)
"In earlier days Wild Asses were fairly common in the Syrian
Desert, and they entered Trans jordania freely. They have since
become rare and are probably on the verge of extinction in the
Syrian desert. Two forms occurred there which correspond to the
two Wild Asses of the Bible, i. e. the Syrian Wild Ass (Equus hemip-
pus) and the Onager (E. onager) ." (Bodenheimer, 1935, p. 116.)
"That the onager was regularly captured and domesticated in
Assyria in ancient times is clearly established by one (Fig. 23) of
the bas-reliefs discovered by Sir A. H. Layard at Kouyunjik
(Nineveh). The relief, which is one of a series of slabs recording
scenes in the life and hunting expeditions of Assur-Bani-Pal (B. C.
668-626), represents two of the king's attendants lassoing a wild
ass. The other asses are seen running away." (Ridgeway, 1905,
p. 48.)
Xenophon (Anabasis, book 1, ch. 5) mentions seeing large numbers
of Wild Asses, in company with Ostriches, in the vicinity of the
Euphrates in 401 B. C.
Porter (1821, pp. 460-461) gives the following brief account of
the animal in Mesopotamia: "I was informed by the mehmandar,
who had been in the desert, . . . that the wild ass of Irak Arabi
[in the lower Tigris-Euphrates Basin] differs in nothing from the
one I had just seen [in the Province of Fars, Persia]. He had
observed them often, for a short time, in the possession of the Arabs,
who told him the creature was perfectly untameable."
In the 1840's Layard (1850, pp. 265-266) observed a large herd of
Wild Asses in the Sinjar region west of Mosul. He adds that those
mentioned by Xenophon must have been seen in these very plains.
"The Arabs sometimes catch the foals during the spring, and bring
them up with milk in their tents. . . . They are of a light fawn
color almost pink. The Arabs still eat their flesh."
"Wild ass ... range over the plain between the Tigris and the
Euphrates, but do not, as generally stated, extend into the Syrian
Desert. The only locality which I know of as being a sure place to
come in contact with these very elusive beasts is the Jebel Sinjar,
13
370 EXTINCT AND VANISHING MAMMALS
between Deir on the Euphrates and Mosul on the Tigris." (Carru-
thers, 1915, p. 22.)
A later account by Carruthers (1935, pp. 147-149) is as follows:
The Syrian Wild Ass . . . had a wide range over the Syrian Hammad in
the 16th and 17th centuries. John Eldred saw wild asses between Hit and
Aleppo in 1584, Cartwright in 1603 beheld "every day great droves of wild
beasts, as wild asses all white," this was not far from Ana on the Euphrates.
Teixeira a few years later saw many herds in the region of Ur in the
Chaldees, while Delia Valle described a captive "wild ass or little onager"
which he saw in the piazza before the Pasha's house in Basra in 1625 ....
It would appear that the Wild Ass disappeared from the Syrian Desert
during the 18th century, and was exterminated in Northern Arabia during
the 19th. Burckhardt reported that they were still numerous in the Shararat
country in the first decade of last century. Its last refuge appears to have
been in the lava country to the south-east of Jabal Druz. Musil says [1927
or 1931] "I have heard that as late as a hundred years ago there were Wild
Asses roaming near the depression of Sirhan, where they had an abundance
of water and, in the volcanic district, good pasture and still better hiding
places. It is said that the last Wild Ass was shot at the wells of Al Ghamr,
[34 miles] south-east of the lake of Azrak. Old Hmar told stories of his
grandfather's hunts for Wild Asses near the depression of Sirhan; but since
firearms have come to be used by the Bedouins, Wild Asses have become
less and less numerous. They are still to be found in the Jezire, between the
middle Euphrates and Tigris, whence the Sleyb often bring their Asses for
breeding purposes." Guarmani confirms this custom of crossing the domestic
asses with the wild ones, and also the fact of the extermination of the latter
south of the Euphrates. He says "When winter comes, many of the Saleib
cross the Euphrates to hunt the Wild Ass in Mesopotamia, there being no
more of these now (1865) in the Hammad. They take a certain number
of them alive to breed with their own." . . .
It is almost certain that they have now been exterminated in their last
refuge, north of the Euphrates, the Jabal Sin jar, none having been seen
since 1927. Whether or not there are a few left in South Arabia, or in the
Oman hinterland, seems doubtful.
The British Museum has a specimen from Mesopotamia, presented
by Layard before 1852, and a Syrian specimen, received from the
Zoological Society of London in 1867. No modern zoologist seems
to have met with this subspecies in the field, and wild-killed animals
are evidently among the rarest of all museum specimens.
Antonius (1928, pp. 19-20) records a male that had been received
in 1911 from the "desert north of Aleppo," Syria, and was still
living in the Schonbrunn Zoo in 1928; also three preserved specimens
that had lived at Schonbrunn in the latter part of the past century.
"The little Hemippus ... of Mesopotamia and Syria, domesti-
cated by the ancient Sumers before the introduction of the horse,
. . . became perhaps totally extinct in recent years. It could not
resist the power of the modern guns in the hands of the Anazeh and
Shammar nomads, and its speed, great as it may have been, was not
sufficient always to escape from the velocity of the modern motor
ORDER PERISSODACTYLA I ODD-TOED UNGULATES 371
car which more and more is replacing the Old Testament Camel-
Caravan." (Antonius, 1938, p. 559) .
In a later paper (1939) Antonius makes out a strong case for the
domestication of the Syrian Wild Ass by the Sumerians in the third
millennium B. C.
Shooting of gazelles and other game from motor cars in now said
to go on throughout the Syrian and Arabian Deserts (J. C. Phillips,
in litt., June, 1938). Perhaps this modern "sport" was instituted
before the last Syrian Wild Asses had been killed, and it may have
been the final factor in their disappearance.
The Onager of Anatolia
ASINUS HBMIONUS SUbsp.
Of this animal there evidently remains nothing but a tradition.
It was presumably a subspecies of Asinus hemionus; but whether it
was the North Persian onager, the Syrian hemippus, or some unde-
scribed form, we shall probably never know. It must have inhabited
the rolling downs that Carruthers describes (1915, p. 10) as the
habitat of the Anatolian Wild Sheep (Ovis ophion anatolica).
Pliny (Hist. Nat., VIII, 44) reported "Onagers" in ancient Phrygia
and Lycaonia, corresponding more or less to the modern Anatolia.
"In early days the wild ass was well known in Paphlagonia
[a country on the south of the Black Sea] , for Homer, when speaking
of the Eneti who came from thence to aid Priam and the Trojans,
describes their land as 'the home of wild mules.' There can be little
doubt that the wild mule of Paphlagonia was some form of Equus
hemionus, probably the same variety as that called 'mule' (he-
mionus) in Aristotle's time." (Ridgeway, 1905, pp. 50-51.)
"The Onager of Anatolia, so well known to Pliny and other an-
cient authors, was exterminated before modern times" (Antonius,
1938, p. 559).
Family TAPIRIDAE: Tapirs
This family is represented by two genera (Tapirus and Tapirella)
in Central and South America and by one genus (Acrocodia) in
southeastern Asia and Sumatra. Of the seven New World forms,
ranging from Mexico to Argentina, one, Tapirus roulinii, is dealt
with in Dr. Allen's volume. An account of the single Old World
species follows here.
372 EXTINCT AND VANISHING MAMMALS
Malay Tapir
ACROCODIA INDICA (Desmarest)
Tapirus indicus "Cuvier" Desmarest, Nouv. Diet. Hist. Nat., vol. 32, p. 458,
1819. (Malay Peninsula.)
FIGS.: Geoffroy and Cuvier, Hist. Nat. Mammif., pi. 303, 1825; Gervais,
Hist. Nat. Mammif., pt. 2, pi. 51, 1855; Royal Nat. Hist., vol. 2, p. 458,
fig., 1894; Proc. Zool. Soc. London 1908, p. 786, fig.; Kerr, 1927, pi. 7;
Jour. Bombay Nat. Hist. Soc., vol. 37, no. 1, suppl., pi. 34, 1934; Pocock,
1937, p. 710, fig.
It is generally agreed that this interesting animal of the Malay
Peninsula and Archipelago deserves and requires protection, though
its numbers may not yet be reduced to the danger point.
The general form is heavy; limbs short and stout; tail short;
ears oval; eyes small; nose and upper lip produced into a short
proboscis; front feet four-toed; hind feet three-toed; head, limbs,
and front part of body brownish black; body behind the shoulders,
including rump and upper part of thighs, and ear tips grayish white.
Height at shoulder, 36 to 42 inches. Young brownish black, spotted
and streaked with brownish yellow and white. (Blanford, 1891,
pp. 478-479.)
Malay Peninsula and adjacent region. Blanford (1891, p. 479)
reported the Tapir's northern limit at about lat. 15 N. in Tenas-
serim, but Arthur S. Vernay extended the range about 3 farther
north, on the Burmo-Siamese frontier (Jour. Nat. Hist. Soc. Siam,
vol. 6, p. 318, 1924).
Its status in Burma is discussed by Peacock (1933, pp. 201-202) :
Tapirs are found only in the Tenasserim forests in southernmost Burma.
They are found only in very dense evergreen forests, but are much more
common than their retiring habits would lead one to believe. I found tapirs
to be very common indeed along the Big Tenasserim River and in the
Victoria Point Range in the Mergui Forest Division. Tapirs are generally
found in pairs or solitary. . . .
The tapir appears to be singularly blessed in that neither man nor the
carnivora appear to be particularly keen on hunting it. ...
There can be no pleasure or object in shooting a gentle, shy animal that
does not bear even an insignificant trophy. I have heard of tapir being
shot only on two occasions.
Tapirs are wholly protected animals under existing game laws.
"The distribution of the Tapir in Siam is very imperfectly known,
but it seems to occur in Peninsular and South-western Siam. Said
to be fairly common in Patani, and recorded from Hat Sanuk and
Hue Sai near the Siam-Tenasserim frontier." (Gyldenstolpe, 1919,
p. 170.)
Gairdner (1915, p. 141) reports finding tracks of the Tapir in the
Petchaburi Valley, Siam, and adds: "They are never, I believe,
intentionally shot by jungle folk, who look upon these rather
ORDER PERISSODACTYLA I ODD-TOED UNGULATES
373
defenceless creatures as peculiar. They allege that the Creator,
having devised all other beasts to his entire satisfaction, had left
over numerous remnants of clay. Taking these in his hands, he
rolled them all up together .... Hence the tapir."
According to Giles (1936, pp. 167-168), "They are also found in
the forests of Tavoy and Mergui in Burma and in the Malay States
.... The flesh of this animal is not much esteemed as food nor is
the animal sought after for commercial purposes. The reason for
this animal having survived may be found in these two main factors.
The chief enemy of the tapir is the tiger."
FIG. 39. Malay Tapir (Acrocodia indica)
In 1933 total protection was recommended by the Siam Society.
In Cambodia Tapirs have almost completely disappeared at the
present time (Resident Superieur of Cambodia, in Hit., November 20,
1936).
In southeastern Indo-China the species was still fairly common
15 years ago, but it is becoming rarer and rarer. It was believed to
have practically disappeared, when last year a report came from a
trustworthy source that two had been killed by native trappers in
the Hongquan district of eastern Cochin China. The species is
absolutely protected under the law. (Andre Kieffer, in Hit., Novem-
ber 21, 1936.)
In the Malay Peninsula, according to Ridley (1895, pp. 161-162),
"This animal is still tolerably abundant in the further jungles of the
interior, and though rarer than the rhinoceros, is oftener to be seen
in captivity. . . . When taken young it becomes very tame and
374 EXTINCT AND VANISHING MAMMALS
amusing. As the jungles are cleared the tapir becomes rarer and
rarer."
S. S. Flower (1900, pp. 368-369) gives records for Province Welles-
ley, Kedah, Malacca, Pahang, and Perak, and adds: "H. H. the
Rajah Mudah of Kedah told me (June 1898) that the Tapir is not
uncommon in the swamps of Kedah, within a day's journey of Alor
Star. Mr. F. H. Malcolm Staples told me (Sept. 1897) that the Tapir
is still sometimes met with about Batu Pahat, Johore."
It is "still not uncommon in the Malay Peninsula and Sumatra,
but much persecuted by menagerie keepers. No zoo anywhere in the
world is considered complete without a pair of Malayan tapirs and
for every animal that survives in an exhibition, several die in Singa-
pore, or in transit." It needs protection. (F. N. Chasen, in litt.,
March 31 and May 5, 1937.)
Sumatra. "Concerning the possibilities of the survival of the
Indian Tapir opinions are at variance. Some speak of 'relentless
hunting' by Natives, while others claim that the tapir needs no
protection by law, since it has no market value, the meat is not eaten,
and the animal is, moreover, very shy. In addition, owing to Native
superstition it is little molested. . . . They are yet quite numerous
near the headwaters of the Koealoe and Bila Rivers; as well as
farther south near the kotta Siak Sri Indrapoera and the Mandau
River; as well as near the salt springs of Ampoe Gadang, Djambi
and N. West Palembang.
"Few will question the necessity of enforcing rigidly and thor-
oughly the Decree on Game Protection with regard to this animal."
(Heynsius-Viruly and Van Heurn, 1936, p. 50.)
"I know of one authentic case near the borders of our plantation
[at Dolok Merangir, east-central Sumatra] in which a group of
native clerks were hunting at night. They saw a pair of eyes and
shot to find that they had a large tapir ; one of a pair known to be
in that region. It seems that rhinoceros and other large animals are
killed by the same methods and by planned attacks by the natives
without any serious consequences." (Walter N. Bangham, in litt.,
1933.)
According to Dammerman (in Skottsberg, 1934, p. 422), the
species is threatened with extinction.
Kuiper (1926, pp. 425-426) records three black individuals from
the Palembang region, southeastern Sumatra. He considers that
these represent "more than an individual aberration," and gives
them the name Tapirus indicics var. brevetianus.
"The Malay tapir is strictly protected by Dutch law in Sumatra ;
not even scientific institutions being allowed to collect it. ...
"Carl Berthold, the well-known animal dealer of Medan, has noted
ORDER PERISSODACTYLA : ODD-TOED UNGULATES 375
that tapirs in the wilds suffer from an eye disease and are often
blind." (Ulmer, in Miller, 1942, p. 161.)
Various authors, including De Beaufort (1926, p. 61), extend the
Malay Tapir's range to Borneo, while others consider it confined to
Sumatra within the Malay Archipelago.
"It is not yet certain that the tapir has been met with in Borneo,
although there are persistent reports that an animal of its size and
appearance exists in the interior of the country. It would be wise to
suspend our judgment for the present and content ourselves with the
fact that so far it has only made its appearance on the North
Bornean postage stamps!" (Mjoberg, 1930, p. 22.)
Family RHINOCEROTIDAE: Rhinoceroses
Two genera (Ceratotherium and Diceros) , of two forms each,
occur in southern and eastern Africa, and from the Sudan westward
to Nigeria. Two additional genera (Rhinoceros and Dicerorhinus) ,
consisting of four or five forms, range from India and Indo-China
through the Malay Peninsula to Sumatra and Borneo. Unfortu-
nately, a work of the present scope requires a discussion of every
living form of rhinoceros.
Great Indian Rhinoceros; Great One-horned Rhinoceros.
Rhinoceros unicorne (Fr.)
RHINOCEROS UNICORNIS Linnaeus
[Rhinoceros} unicornis Linnaeus, Syst. Nat., ed. 10, vol. 1, p. 56, 1758. ("Habitat
in Africa, India" (Linnaeus) ; "probably the sub-Himalayan Tarai of
Assam" (Lydekker, 1916, vol. 5, p. 48).)
FIGS.: Geoffrey and Cuvier, 1824, vol. 2, pis. 306, 307; Gervais, Hist. Nat.
Mammif., pt. 2, pi. facing p. 164, 1855; P. L. Sclater, 1876, pi. 95; Royal
Nat. Hist., vol. 2, pi. facing p. 464, 1894; Lydekker, 1900, pi. 1, fig. 2;
Van der Byl, 1915, pi. 32; Faunthorpe, 1924, pp. 174, 181, figs.; New York
Zool. Soc. Bull., vol. 27, p. 72, fig., 1924; Jour. Bombay Nat, Hist. Soc.,
vol. 37, no. 1, suppl., pi. 31, 1934; Pocock, 1937, p. 709, fig.
With a former range extending from the North-West Frontier
Province of India eastward perhaps as far as French Indo-Qiina,
this species has more recently become restricted largely or wholly
to the Nepal Terai, northern Bengal, and Assam. Its numbers also
have greatly diminished.
This is the largest of the Asiatic rhinoceroses, reaching a height
of 6 feet 4 inches at the shoulder and a total length of 14 feet 1 inch,
with a horn of 24 inches; fold of skin in front of shoulder not con-
tinued across back of neck; other folds behind shoulder, in front of
and across thigh, and around the neck; sides of body and upper
limbs studded with large rounded tubercles; skin naked except for
a fringe of hairs on the margin of the ears and some bristly hairs on
376
EXTINCT AND VANISHING MAMMALS
the tail; general color uniformly blackish gray, with more or less
pink on the margins of the folds (Lydekker, 1900, pp. 21-22) .
India. "In the history of Timur-bec, it is described how in 1398
on the frontier of Kashmir, Timur hunted and killed many rhi-
noceroses. In the memoirs of Baber it is described how in about
1519 he hunted the rhinoceros in bush country near the Indus. And
in the book of Sidi Ali dated 1554 it is stated that rhinos were seen
near the Kotal Pass, west of Peshawar.
FIG. 40. Great Indian Rhinoceros (Rhinoceros unicornis)
"These references are of interest, for they show that in old times
the rhinoceros was plentiful and further, ranged over a great portion
of India, whereas it is now approaching extinction." (Hobley, 1931,
p. 19.)
"Not improbably . . . the rhinoceroses found till about the year
1850 in the grass-jungles of the Rajmehal Hills, in Bengal, belonged
to the present species. Now, however, this huge animal has re-
treated almost, if not entirely, to the eastward of the Tista valley,
on the borders of Kuch Behar; its main strongholds being the great
grass-jungles of that province and of Assam." (Lydekker, 1900,
p. 23.)
Shebbeare (1935, pp. 1229-1231) gives the following account:
Though this rhinoceros is becoming alarmingly rare everywhere, Nepal and
Assam are better off than Bengal, where its habitat is restricted to a few
places in the Duars and Cooch Behar State. Here the last main stronghold
of the species is a tract of high grass savannah along the Torsa river,
stretching from the foothills of Bhutan, through the Duars into Cooch Behar.
It is a narrow strip, not more than 40 miles from the north to the south
and, at its widest, four miles from east to west perhaps 50 or 60 square
miles. Outside this tract the few scattered colonies can perhaps muster a
ORDER PERISSODACTYLA: ODD-TOED UNGULATES 377
dozen individuals in all, but unfortunately these outliers have no spare
coverts into which they can expand. . . .
Contrary to what one hears of African rhino, ours is seldom aggressive,
nor does he cause havoc to agricultural crops like the elephant.
For the last 25 years in Bengal and Assam rhino have been closed to
sportsmen, but this has not saved them from poachers, who shoot them to
obtain their horns. From time immemorial these have been highly prized
for superstitious reasons. A cup made of the horn of a rhinoceros is still
believed to render poison innocuous, a point of some importance to tyran-
nical rulers, and, when powdered, it is held in the East, especially in China,
to be the most potent aphrodisiac. It is believed that most of the horns
that are smuggled out of these jungles eventually find their way to China,
but however this may be their present value in the Calcutta market is about
half their weight in gold. A single horn retrieved from the poachers recently
fetched 150 pounds, and still higher prices have been known. That an
animal by nature condemned to carry such a price on his nose should tempt
poachers is not to be wondered at, but the remoteness of their strongholds,
and their armour, too thick to be penetrated by "gas-pipe" guns, was their
protection, and up to about six years ago there were probably some 200
animals living in the small tract I have described.
Then poaching began. The first poachers came from Assam, where they
had plied the same trade, and brought with them muzzle-loading guns heavy
enough to kill a rhino. They were j pined by local men of the same tribe
(Mechs) and formed themselves into gangs. Their plan was to build a
light bamboo staging about 8 ft. above the ground at strategic points, usually
where two well-worn rhino tracks met, and lie up when ths moon was nearly
full. Sooner or later a victim was bound to pass and received a heavy bullet
at a range of a few feet .... They seldom took more than the horn; to
try to dispose of the meat, which, by the way, is -excellent eating, would
have aroused suspicion .... For nearly three years this went on without
any suspicion being aroused.
After the poaching was detected, it required six months or more of
effort by the Forest Department and the Government of Bengal to
stop the poaching. A bill was passed, making the killing of rhino,
except in defense of life, an offense.
"Our attempts have so far been successful. Since Christmas, 1931,
so far as we know, only one rhino has been killed, and the perpe-
trators are now in gaol."
The Government of Bihar (in litt., December, 1936) sends the
following information: "The Great One-horned Rhinoceros was
formerly fairly common in the jungles of North Bihar bordering on
Nepal, especially . . . near the Kosi river, and individuals were
found until 50 or 60 years ago. The jungles in this area have prac-
tically disappeared and the animal is unknown except as an occa-
sional stray visitor from Nepal into the jungles in the North West
corner of the Champaran district."
The Senior Conservator of Forests, Bengal, writes (in litt., Sep-
tember, 1937) :
"Former range: Jalpaiguri Forests (common) and Riparian
Forests of the Buxa Division (no information as to number) .
378 EXTINCT AND VANISHING MAMMALS
"Present range: 4 or 5 in the Jalpaiguri Forests, 56 in the Riparian
Forests bordering the Torsa and Malangi Rivers and wet forests of
Kanbari.
"Causes of depletion: in Jalpaiguri Forests: probably shooting
and poaching and possibly disease. In the Buxa Division the species
showed up to 1932 tendency to extinction due to heavy poaching.
Since 1933 they are increasing in number. The horn is worth 8 to 10
times its weight in silver. The hide is also valuable. ... A Game
Sanctuary to the extent of 26 sq. miles is being maintained."
In years gone by, in this general region, the animals were suffi-
ciently numerous or destructive to have called for the establishment
of a bounty. "They sometimes will travel long distances to reach
rice and corn fields, and do immense mischief, so much so that
there is a Government reward of twenty rupees to anyone shooting
a rhinoceros" (Baldwin, 1876, p. 144) .
Nepal. This species is "decreasing rapidly in Nepal. In the
Morang District of the Nepal Tarai this rhinoceros was plentiful
not many years ago, but now not a single specimen is, I believe, to be
found within two hundred miles." Several specimens were collected
in the Gandak Valley in 1923. (Faunthorpe, 1924, pp. 179-188.)
Further information is as follows (Anonymous, 1934, p. 89) :
Along the numerous rivers which flow through the jungles of the Nepal
Terai the rhino has particular places for dropping its excreta. Mounds so
accumulate in places. In approaching these spots a rhinoceros walks back-
wards and falls an easy victim to poachers. . . .
The food consists chiefly of grass. In Nepal during the rains Rhinoceros
frequently enter cultivation. . .
In Nepal the flesh and the blood of the Rhinoceros is considered highly
acceptable to the Manes. High caste Hindus and most Gurkhas offer libation
of the animal's blood after entering its disembowelled body. On ordinary
Sradh days the libation of water and milk is poured from a cup carved from
its horn. The urine is considered antiseptic and is hung in a vessel at the
principal